Soil carbon sequestration by three legume pastures is greater in deep soil than in surface soil

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Roots and Associated Fungi Drive Long-Term Carbon Sequestration in Boreal Forest

Roots and Associated Fungi Drive Long-Term Carbon Sequestration in Boreal Forest

Boreal forest soils function as a terrestrial net sink in the global carbon cycle. 北方森林土壤在陆地网络的碳循环网络中发挥着重要作用The prevailing dogma has focused on aboveground plant litter as a principal source of soil organic matter.目前普遍认为地上部分的植物残体是土壤有机物的主要来源Using 14C bomb-carbon modeling, we show that 50 to 70% of stored carbon in a chronosequence of boreal forested islands derives from roots and root-associated microorganisms.通过碳14炸弹碳模型,我们在发现北方森林的土壤年代序列中有50—70%的固定碳源自树根及菌根微生物Fungal biomarkers indicate impaired degradation and preservation of fungal residues in late successional forests. 真菌的生物标记表明了森林演替中真菌残留物的腐败降解和保存Furthermore, 454 pyrosequencing of molecular barcodes, in conjunction with stable isotope analyses, highlights root-associated fungi as important regulators of ecosystem carbon dynamics. 此外,454焦磷酸测序的分子条形码协同稳定同位素分析表明菌根菌是生态系统碳动力学的重要调节者Our results suggest an alternative mechanism for the accumulation of organic matter in boreal forests during succession in the long-term absence of disturbance.本研究认为在长期没有干扰的北方森林的有机物积累演变过程存在一个替代机制。

The knowns, known unknowns and unknowns of sequestration of soil organic

The knowns, known unknowns and unknowns of sequestration of soil organic

Agriculture,Ecosystems and Environment 164 (2013) 80–99Contents lists available at SciVerse ScienceDirectAgriculture,Ecosystems andEnvironmentj o u r n a l h o m e p a g e :w w w.e l s e v i e r.c o m /l o c a t e /a g eeReviewThe knowns,known unknowns and unknowns of sequestration of soil organic carbonUta Stockmann a ,∗,Mark A.Adams a ,John W.Crawford a ,Damien J.Field a ,Nilusha Henakaarchchi a ,Meaghan Jenkins a ,Budiman Minasny a ,Alex B.McBratney a ,Vivien de Remy de Courcelles a ,Kanika Singh a ,Ichsani Wheeler a ,Lynette Abbott b ,Denis A.Angers c ,Jeffrey Baldock d ,Michael Bird e ,Philip C.Brookes f ,Claire Chenu g ,Julie D.Jastrow h ,Rattan Lal i ,Johannes Lehmann j ,Anthony G.O’Donnell k ,William J.Parton l ,David Whitehead m ,Michael Zimmermann naFaculty of Agriculture and Environment,The University of Sydney,Biomedical Building C81,Suite 401,1Central Avenue,Australian Technology Park,Eveleigh,NSW 2015,Australia bSchool of Earth and Environment and UWA Institute of Agriculture M082,The University of Western Australia,35Stirling Highway,Crawley,The University of Western Australia,WA 6009,Australia cAgriculture and Agri-Food,2560Hochelaga Boulevard,Quebec City,Quebec,G1V 2J3,Canada dCSIRO Land and Water and Sustainable Agriculture Flagship,PMB 2,Glen Osmond,SA,5064,Australia eSchool of Earth and Environmental Science and Centre for Tropical Environmental and Sustainability Science,James Cook University,PO Box 6811,Cairns 4870,Australia fRothamsted Research,Harpenden,Herts,AL52JQ,UK gAgroTechParis,France hBiosciences Division Argonne National Laboratory,USA,9700S.Cass Avenue,Argonne,IL,60439,USA iSchool of Environment and Natural Resources,The Ohio State University,USA,422B Kottman Hall,2021Coffey Road,The Ohio State University,Columbus,OH,43210,USA jCornell University,909Bradfield Hall,Cornell University,Ithaca,NY,14853,USA kFaculties of Science,The University of Western Australia,35Stirling Highway,Crawley,The University of Western Australia,WA,6009,Australia lNatural Resource Ecology Laboratory,Colorado State University,NESB B233,Campus Mail 1499,Fort Collins,Colorado State University,CO,80523-1499,United States mNew Zealand Agricultural Greenhouse Gas Research Centre,New Zealand Landcare Research,PO Box 40,Lincoln 7640,New Zealand nUniversity of Natural Resources and Life Sciences Vienna,Department of Forest and Soil Sciences,Institute of Soil Research,Peter Jordan Str.82,A-1190,Wien,Austriaa r t i c l ei n f oArticle history:Received 1August 2011Received in revised form 25September 2012Accepted 2October 2012Available online 23 November 2012Keywords:Soil carbon sequestration Soil carbon poolsSoil carbon modellinga b s t r a c tSoil contains approximately 2344Gt (1gigaton =1billion tonnes)of organic carbon globally and is the largest terrestrial pool of organic carbon.Small changes in the soil organic carbon stock could result in significant impacts on the atmospheric carbon concentration.The fluxes of soil organic carbon vary in response to a host of potential environmental and anthropogenic driving factors.Scientists world-wide are contemplating questions such as:‘What is the average net change in soil organic carbon due to environmental conditions or management practices?’,‘How can soil organic carbon sequestration be enhanced to achieve some mitigation of atmospheric carbon dioxide?’and ‘Will this secure soil quality?’.These questions are far reaching,because maintaining and improving the world’s soil resource is imper-ative to providing sufficient food and fibre to a growing population.Additional challenges are expected through climate change and its potential to increase food shortages.This review highlights knowledge of the amount of carbon stored in soils globally,and the potential for carbon sequestration in soil.It also discusses successful methods and models used to determine and estimate carbon pools and fluxes.This knowledge and technology underpins decisions to protect the soil resource.© 2012 Elsevier B.V. All rights reserved.∗Corresponding author.Tel.:+61286271147;fax:+6193513706.E-mail addresses:uta.stockmann@.au (U.Stockmann),mark.adams@.au (M.A.Adams),john.crawford@.au (J.W.Crawford),damien.field@.au (D.J.Field),mhen8672@.au (N.Henakaarchchi),meaghan.jenkins@.au (M.Jenkins),budiman.minasny@.au (B.Minasny),Alex.McBratney@.au (A.B.McBratney),vivien.deremydecourcelles@.au (V.d.R.d.Courcelles),kanika.singh@.au (K.Singh),ichsani.wheeler@.au (I.Wheeler),lynette.abbott@.au (L.Abbott),denis.angers@agr.gc.ca (D.A.Angers),Jeff.Baldock@csiro.au (J.Baldock),michael.bird@.au (M.Bird),philip.brookes@ (P.C.Brookes),Claire.Chenu@grignon.inra.fr (C.Chenu),jdjastrow@ (J.D.Jastrow),lal.1@ (l),CL273@ (J.Lehmann),tony.odonnell@.au (A.G.O’Donnell),billp@ (W.J.Parton),WhiteheadD@ (D.Whitehead),michael.zimmermann@boku.ac.at (M.Zimmermann).0167-8809/$–see front matter © 2012 Elsevier B.V. All rights reserved./10.1016/j.agee.2012.10.001U.Stockmann et al./Agriculture,Ecosystems and Environment164 (2013) 80–9981 Contents1.Introduction (81)1.1.What is the overall purpose of soil carbon sequestration? (81)1.2.How do we support SCS as a science community? (81)2.Soil—A terrestrial pool of organic carbon (82)2.1.Effects of climatic conditions and ecosystem conditions on SOC pools (82)2.1.1.Climatic conditions (82)2.1.2.Ecosystem conditions and land use change (82)2.2.Environmental conditions and decomposition (82)2.3.Biological processes affecting the decomposition of SOC (83)2.3.1.Priming effects (83)2.3.2.Biodiversity (84)2.3.3.Roots and root exudates (84)2.4.Chemical and physical processes affecting the decomposition of SOC (85)2.4.1.Does the solubility of SOM,as determined by chemical(alkaline/acid)extractions,sufficiently characterizerecalcitrant SOM? (85)2.4.2.Is distinction of SOM in terms of soil function and as conceptualized in process models of SOM dynamics,driven byknowledge of rates of decomposition and humification pathways? (85)2.4.3.Can the physical nature of SOM be better represented by a‘molecular aggregate’model as proposed byWershaw(2004)in place of a‘humic polymer’model? (86)2.4.4.Taken together,these questions suggest a new view of decomposition processes and recalcitrance (86)3.Soil carbon measurement—Pools andfluxes (86)3.1.SOC pools (86)3.1.1.Measurement of SOC (86)3.1.2.SOC fractionation (86)3.1.3.Imaging SOC in situ with structure (86)3.2.Cfluxes and their measurement (87)4.Soil carbon modelling (88)4.1.Process-oriented versus organism-oriented models (88)4.1.1.Model characteristics (88)4.1.2.Model limitations (90)4.2.Other models used and currently being developed to potentially predict SOM stocks orfluxes (92)4.2.1.Empirical regression models (92)ndscape models (92)4.2.3.‘Whole systems’modelling (92)4.2.4.Prospects for potential‘Next Generation’soil carbon models (92)4.3.Where to from here in modelling of SOM dynamics? (93)5.Practical measures for enhancing soil carbon (94)6.Conclusions (95)Acknowledgements (96)References (96)1.IntroductionApproximately8.7Gt(1gigaton=1billion tonnes)of carbon(C) are emitted to the atmosphere each year on a global scale by anthro-pogenic sources(Denman et al.,2007;Lal,2008a,b).However,the atmospheric increase has been in the order of3.8Gt C yr−1(rate of increase in the year2005,Denman et al.,2007),highlighting the important regulatory capacity of biospheric C pools(Le Quéréet al., 2009).In this context,soil organic carbon(SOC)and its potential to become a‘managed’sink for atmospheric carbon dioxide(CO2)has been widely discussed in the scientific literature(e.g.Kirschbaum, 2000;Post and Kwon,2000;Guo and Gifford,2002;Lal,2004a,b, 2008a,b;Post et al.,2004,2009;Smith,2008;Chabbi and Rumpel, 2009;Luo et al.,2010b).Here,we use the term SOC to define C in soil derived from organic origins.The term soil organic matter(SOM) is also used frequently in the literature and is generally agreed to contain about58%SOC(i.e.elemental C).SOM is a mixture of mate-rials including particulate organics,humus and charcoal along with living microbial biomass andfine plant roots.To reward‘good’management of the soil C pool leading to enhanced soil carbon sequestration(SCS),there are a number of overarching questions that need to be considered in relation to the potential of the soil–plant system to‘sequester’organic C,where sequestering soil carbon requires a stipulated duration timeframe (usually100years)in order to be considered a‘permanent’increase under managed agricultural systems.SCS implies an increase in soil C for a defined period against a baseline condition where the increased C is sourced from atmospheric CO2.This implication helps to frame the following questions:1.1.What is the overall purpose of soil carbon sequestration?What is the value in increasing the inputs to soil organic matter aside from its role in potential sequestration?For how long must this increase be maintained to be considered as SCS?Consequently, what is more important,long-term SCS or the functioning of the soil?Are these roles essentially inseparable?1.2.How do we support SCS as a science community?How can the benefits of SCS be promoted among policy makers/farmers/landholders(i.e.the potential of SCS to mitigate climate change,the use of SCS as a platform for sustainable agriculture)and how can suitable answers to questions such as measurement,modelling,monitoring and permanence for SCS and/or management advice be followed through?Is there a need to improve models of SOM dynamics in order(i)to demonstrate bet-ter understanding of the functioning of the soil ecosystem and(ii) to better assist landholders/farmers with management decisions?82U.Stockmann et al./Agriculture,Ecosystems and Environment164 (2013) 80–99Can both of these questions be addressed within the same model?This article synthesizes current soil C research and highlights a number of key research areas.2.Soil—A terrestrial pool of organic carbonGlobally,the quantity of C stored in the soil is second only to that in the ocean(38,400Gt).While the terrestrial biotic C pool is ∼560Gt of organic C(Fig.1),the soil C pool is more than four times thisfigure.The organic C pool capacity of world soils has been vari-ously estimated for principal biomes(refer to Table1).For instance, approximately2344Gt of organic C is stored in the top three meters of soil,with54%or1500Gt of organic C stored in thefirst meter of soil and about615Gt stored in the top20cm(Jobbágy and Jackson, 2000;Guo and Gifford,2002).This contrasts with the∼9Gt addition of anthropogenically liberated‘new’C that is added to the atmo-sphere annually from fossil C sources(coal,oil and gas)and through ecosystem degradation.Several points quickly follow.For exam-ple,a change of just10%in the SOC pool would be equivalent to 30years of anthropogenic emissions and could dramatically affect concentrations of atmospheric CO2(Kirschbaum,2000).Alterna-tively,small increases in rates of oxidation of soil C as a result of increasing temperatures could result in further increases in atmo-spheric CO2(Davidson and Janssens,2006).In general,plant production and patterns of biomass alloca-tion strongly influence relative distributions of C with soil depth (Jobbágy and Jackson,2000).The deeper in the soil profile,the older stored SOC is likely to be.For example,Trumbore(2009)postu-lated that low-density C and microbial phospholipid acids would increase in age with soil depth.Fontaine et al.(2007)proposed an increase of mean residence times of SOC of up to2000–10,000 years for depths beyond20cm.Increased mean residence times reflect reduced microbial activity and SOC turnover at depth.This conceptual model is supported by patterns of root biomass and rel-ative root density that also decline with soil depth(Jobbágy and Jackson,2000)and by increasing concentrations of organo-mineral complexes with depth(Fontaine et al.,2007).2.1.Effects of climatic conditions and ecosystem conditions onSOC pools2.1.1.Climatic conditionsWorldwide,SOC stocks generally increase as mean annual tem-perature decreases(Post et al.,1982).Cool/cold,humid climate regions are characterized by their C-rich soils(Hobbie et al.,2000); for example,approximately1672Gt of C is stored in the arctic and boreal ecosystems of the northern hemisphere—a large proportion of the world’s soil C(Tarnocai et al.,2009).Increased greenhouse gas concentrations in the atmosphere are set to accelerate the rate of warming that may in turn change net primary productivity(NPP),the type of organic matter inputs to soil and soil microbial activity are all important drivers of SOCfluxes. This may release additional CO2from some soils.Kirschbaum (2000)concluded that global warming is likely to reduce SOC by stimulating rates of decomposition whilst simultaneously increas-ing SOC through enhanced NPP resulting from increased CO2levels with the net change in SOC stocks expected to be small over the coming centuries.On the other hand,Sitch et al.(2008)propose that in some instances,soil might be a comparatively stronger source of CO2in the future as temperature rises.2.1.2.Ecosystem conditions and land use changeTypically,arable soils contain around1–3%of SOC,whilst grass-land and forest soils usually contain more(Jenkins,1988).Guo and Gifford(2002)highlighted the influence of land use changes on soil C stocks.Their meta-analysis of74publications suggested that land use changes from pasture to plantation(−10%),native forest to plantation(−13%),native forest to crop(−42%)and pasture to crop(−59%)reduced total C stocks whereas changes from native forest to pasture(+8%),crop to pasture(+19%),crop to plantation (+18%)and crop to secondary forest(+53%)increased total C stocks.A reasonable summary is that changing land use from cropland to pasture or cropland to permanent forest result in the greatest gains of SOC.Post and Kwon(2000),for example,estimated aver-age rates of SOC accumulation for pasture and forest establishment to be of0.33t C ha−1yr−1and0.34t C ha−1yr−1,respectively.Con-version from nearly all other land uses to cropping or monocultures result in losses of SOC.The effects of tillage and diversity are clearly important research foci.The quantity,quality and timing of organic matter inputs to soil vary with species composition within community types(i.e. relative abundance of N-fixing species)as well as with whole-sale changes in community structure(i.e.cropland,grassland, shrubland,woodland,forest).Changes may also result from either management or natural variations in edaphic conditions at a local scale(e.g.Binkley and Menyailo,2005;Hart et al.,2005).These interactions can be complex and vary through time.A recent study investigating long-term(1930–2010)dynamics of SOC after land use change in Java,Indonesia(Minasny et al.,2011),showed that after nearly40years of decline as a result of conversion of primary forest to plantation and cultivated land,total SOC increased,begin-ning around1970.This switch,from decreasing to increasing most likely resulted from human interventions to increase plant produc-tion through fertilizer application and other management(i.e.the ‘Green-Revolution’)The total SOC in Java showed a net accumula-tion rate of0.2–0.3t C ha−1yr−1in thefirst10cm during the period of1990–2000.2.2.Environmental conditions and decompositionSOM is derived from the microbial decomposition of plant inputs,either directly,as plant residues,or indirectly,as ani-mal residues.During decomposition,the intact plant and animal residues are initially broken down into small particles of largely intact material.Eventually,following repeated recycling through the soil micro-organisms,as transient products of decomposi-tion,original plant or animal residues give rise to a highly stable, black-brown substance referred to as humus.Globally,humus is calculated to have a mean turnover time of27years(Jenkins,1988).It is now generally considered that SOM decomposition is con-trolled more by biological and environmental conditions than by molecular structures of the carbon-based inputs(Schmidt et al., 2011).SOM is thus described as a continuum of materials in vary-ing states of decomposition,with the chemical composition at any given site dependent upon the interplay of site conditions and bio-logical limitations.Litter provides the C that supports heterotrophic microbial activity,and the more readily decomposable the C,the more rapidly the microbial community can grow(Agren and Bosatta,1996). Decomposition returns to the atmosphere most of the C added in litter to the soil surface—only a very small fraction becomes humus.Through their effects on microbial activity,moisture and temperature exert strong control over the rate of litter decomposi-tion,followed by litter quality(decomposability)and soil microbial community composition(Meentemeyer,1978;Melillo et al.,1982; Parton et al.,2007).Moisture and temperature also exert strong control of humus decomposition.There is currently significant debate in the scientific community about the temperature sensitivity of different frac-tions(e.g.litter vs humus)and different pools(bile vs stable) of organic matter mineralization.At present a single relationshipU.Stockmann et al./Agriculture,Ecosystems and Environment164 (2013) 80–9983Fig.1.Total annualflux of carbon in gigatons(billions of tonnes)through the most biologically active pools(as compared to the deep-ocean and lithosphere).Natural annual input of‘new’C to the biosphere through volcanism and rock dissolution is balanced by long term burial of C in ocean sediments.Annual anthropogenic input of‘new’C into the atmospheric pool(averaged over the last40years)comes from land clearing(1–2Gt annually)and fossil fuel sources(8.7Gt C annually in2008(Le Quéréet al.,2009)). Mean uptake rates of terrestrial and ocean CO2sinks are2.6and2.2Gt C yr−1for1990–2000,respectively(Denman et al.,2007).If not cited otherwise,estimates adapted from Beer et al.(2010),Denman et al.(2007)and Volk(2008).(i.e.a unique Q10),is applied to all SOM pools in most decompo-sition models.On the other hand,chemical theory predicts that recalcitrant forms of organic matter should be more sensitive to temperature than labile forms.In the literature,results vary widely (Giardina and Ryan,2000;Fang et al.,2005;Craine et al.,2010).2.3.Biological processes affecting the decomposition of SOC2.3.1.Priming effectsPriming effects werefirst recorded more than50years ago (Bingeman et al.,1953).In a review Kuzyakov et al.(2000)defined soil priming as“strong short-term changes in the turnover of SOM caused by comparatively moderate treatments of the soil”.‘Prim-ing effects’of added C(or N)on rates of mineralization of SOM are now well documented(e.g.Kuzyakov et al.,2000;Fontaine et al.,2003;Sayer et al.,2007),and may be either positive(i.e. increased C and N mineralization)or negative(i.e.immobilization). Addition of purified labile C substrates such as glucose(Dilly and Zyakun,2008),fructose and oxalic acid(Hamer and Marschner, 2005),or cellulose(Fontaine et al.,2007)generally stimulate min-eralization of SOM.Explanations of the priming effect include co-metabolism—additions of labile or fresh C stimulate growth of a suite of microorganisms that in turn leads to an increase in microbial enzyme production(Kuzyakov et al.,2000;Hamer and Marschner,2005).Fontaine et al.(2003)hypothesized that changes in land use and agricultural practices that increase the distribution of fresh C at depth could stimulate the mineraliza-tion of ancient buried C.For example,a change from shallow to deep-rooted grasses might have an overall negative effect on SOC due to the potential release of C buried at depth.Alternatively, might increased allocation of C below ground outweigh this poten-tial effect?It remains to be seen if all soil microorganisms follow the same laws or if possible differences in inherent characteris-tics affect microbial diversity and SOC sequestration(Chabbi and Rumpel,2009).Quantifying priming effects underfield conditions is challeng-ing.Isotopic labelling is a preferred approach and well suited to small scale laboratory experiments,but difficult to apply in the field.In the laboratory,priming effects are generally of short dura-tion and of small monly,once the priming effect has dissipated,soil metabolism rapidly reverts to background rates. In systems where SOC accumulates most rapidly(i.e.some grass-lands where a large proportion of NPP is belowground)priming effects may be of significance to total SOC.The true significance of priming effects to SOC on a global,or even ecosystem scale, awaits evaluation.It seems likely that priming effects will remain84U.Stockmann et al./Agriculture,Ecosystems and Environment164 (2013) 80–99Table1Estimates of SOC sink capacity of world soils,listed for principal biomes.Biome SOC storage(Gt C)by soil depth Reference0–1m0–2m0–3mTropical regions354–4031078–1145Batjes(1996)aOther regions616–6401760–1816Global estimate1463–15482376–2456Boreal forest112141150Jobbágy and Jackson(2000)b Cropland157210248Deserts112164208Sclerophyllous scrubs76104124Temperate deciduous forest122145160Temperate evergreen forest7391102Temperate grassland105143172Tropical deciduous forest119175218Tropical evergreen forests316408474Tropical grassland/savannah198281345Tundra114133144Global estimate150219932344Boreal forest338IGBP(Carter and Scholes,2000) Cropland165Deserts and semi deserts159Wetlands N.A.Temperate forest153Temperate grassland/shrubland176Tropical forest213Tropical grassland/savannah247Tundra115Global estimate1567Boreal forest471WBGU(1988)Cropland128Deserts and semi deserts191Wetlands225Temperate forest100Temperate grassland/shrubland295Tropical forest216Tropical grassland/savannah264Tundra121Global estimate2011a Estimates based on4353soil profiles.b Estimates based on2700soil profiles.of greater significance to short-term effects–such as pulses of nutrient availability and of CO2and other greenhouse gases(GHG) to the atmosphere–than to long-term C sequestration.2.3.2.BiodiversityIncreased plant diversity can increase SOC(e.g.Tilman et al., 2006)but this effect,while corroborated by some studies(see Sec-tion2.1.2above),is easier to demonstrate in artificial than‘natural’settings.Many studies of litter decomposition contrast the influences of single plant species,and sometimes mixtures of leaf litter on decomposition(i.e.Melillo et al.,1982;O’Connell,1986;Chapman et al.,1988;Briones and Ineson,1996;Hobbie,1996;McTiernan et al.,1997;Xiang and Bauhus,2007).Much of the influence of indi-vidual plant species on soil microbial activity and nutrient cycling is through the quality and quantity of organic matter returned to the soil(Galicia and García-Oliva,2004;Binkley and Menyailo, 2005).Plant species that host N-fixing bacteria(such as legumes)or root systems with mycorrhizal associations,often enhance nutri-ent uptake and can provide a pathway for the return of C substrate directly to microbes and soil(Hobbie,1992).Increased microbial nitrogen(N)inputs,therefore,most likely increase soil C stocks by influencing decomposition processes.Studies by Kaye et al. (2000)and Resh et al.(2002)showed that compared with Euca-lyptus species N-fixing trees increased soil N availability as well as the potential to sequester SOC.Moreover,N-fixing species tended to add more fresh C to the soil and that C was preferentially decom-posed over older forms of C.2.3.3.Roots and root exudatesRoot-derived organic matter inputs are chemically diverse, ranging in complexity from readily decomposable substrates such as soluble sugars,amino and organic acids,to substrates from root turnover that require greater energy investment to decompose.It has been widely assumed that easily degradable plant exudates are almost exclusively degraded by bacteria(see reviews by Jones, 1998),with fungi playing important roles in the degradation of recalcitrant organic materials such as lignin,as well as cellulose and hemi-cellulose(reviewed by Boer et al.,2005).Perhaps owing to the difficulty(relative to inputs of leaf carbon to soil)of knowing exactly the amounts and chemical nature of root inputs,there is a wide range of views in the literature as to the significance of root inputs to soil carbon stocks andfluxes.Some research suggests root inputs to soil represent5–33%of daily photoassimilate(Jones et al., 2009).Exudates fall into two categories:those that are a result of passive diffusion and over which the plant has little control,and those that have functional significance and at least some degree of regulation of their exudation by plants(see reviews by Jones et al., 2004;Paterson et al.,2007).There are also suggestions,however,that root exudates con-tribute to depletion of SOC stocks through a‘rhizosphere priming effect’based on evidence that overall rates of SOC decompositionU.Stockmann et al./Agriculture,Ecosystems and Environment164 (2013) 80–9985may increase dramatically(up to5-fold)in response to root exu-dates(see Sanderman et al.,2010for examples).Root-C seems to be preferentially stabilized compared to shoot-C(see review by Rasse et al.,2005).Greater chemical recalcitrance of root tissues as compared to that of shoots seems to be responsible for only a small portion of this preferential stabilization.Other SOM protection mechanisms might be enhanced by root activities,such as physico-chemical protection of root exudates,especially in deep soil horizons,and micrometer-scale physical protection through mycorrhiza and root-hair activities that place root C in very small pores and aggregates.This may have implications for C storage in as much as it might suggest;favouring rotations,cropping systems and plant species that allocate C below ground.Much more research is needed to determine and predict the balance between gain versus losses as precipitated by root growth and turnover.2.4.Chemical and physical processes affecting the decompositionof SOCSOC can be protected in the soil matrix through physical-(chemical)stabilization processes,as well as by inherent‘chemical recalcitrance’.Decomposition has been described as a key or even a‘bottleneck’process in C and nutrient cycling(Attiwill and Adams,1993).The chemical composition of decomposing material,especially char-acteristics such as C:N ratios and lignin content,are crucial for determining how quickly decomposition proceeds(Meentemeyer, 1978;Melillo et al.,1982).In general,litter decomposition rate is considered negatively related to C:N ratios,lignin content and lignin:N ratios,and positively related to N concentrations(Melillo et al.,1982,1989).Even so,litter dynamics appear to differ from SOM dynamics.The mineralization of SOM proceeds at a much slower rate than the decomposition of the plant and ani-mal residues from which it is formed.The latter,containing large polymeric molecules of biological origin such as proteins,carbohy-drates,cellulose,etc.are highly favoured for enzymic attack,due to their relatively simple and regularly repeated chemical struc-tures.In contrast,SOM lacks such a simple structure,and is a highly unfavourable substrate for enzymic mineralization(Kemmitt et al., 2008).Recent stable isotope-based research,as synthesized by Amelung et al.(2008),has shown that the residence time of SOM is not correlated to its chemical composition.As a consequence,while relative chemical complexity helps explain short-term decompo-sition of litter and added organic matter decomposition(e.g.at seasonal,annual scales),it does not explain SOM decomposition in the long term(decades,centuries).Mineralization is often directly linked to SOM via soil micro-bial communities and their molecular size,specific activity or composition(Marschner and Kalbitz,2003;Fontaine and Bardot, 2005).A recent,somewhat‘controversial’hypothesis challenges this conventional view and proposes that the mineralization rate of humified SOM is independent of the size,structure or activ-ity of the soil microbial community.Based on observations of fumigation experiments,Kemmitt et al.(2008)argued that the rate-limiting step in SOM mineralization is governed by abiotic rather than microbial processes termed the‘Regulatory Gate’.The‘Regu-latory Gate’hypothesis processes include,among others:diffusion, desorption from soil surfaces,oxidation or stabilized extracellu-lar enzymes.Further research is required to understand better the significance of abiotic versus microbial processes in SOM mineral-ization at the global scale.The regulatory gate hypothesis might suggest controls over C stocks depend on abiotic mechanisms of protection of C and our(human)capacity to influence those. Adsorption/desorption of SOM appears difficult to control in soils, but soil structure is strongly affected by soil management and land use.Certainly,the‘Regulatory Gate’theory is not yet universally accepted(Kuzyakov et al.,2009).Two main physico-chemical)stabilization processes are:(i)pro-tection within aggregates which translates to spatial inaccessibility of soil microbes to organic compounds and a limitation on O2avail-ability and,(ii)interactions with mineral surfaces and metal ions (e.g.see Six et al.,2004;Von Lützow et al.,2006for details).These aspects are of importance because both can provide a priori limits to the soil C sequestration potential of some soils(via the surface properties of minerals such as functional groups and charge)and capacity for soil aggregation and stability(as influenced by soil par-ticle size distribution).It must be noted that increasing SOM has positive effects on both,the aggregation of a soil and the amount of surface charge present.Ongoing discussions regarding the chem-ical and physical processes determining the composition of SOM were reviewed in detail by Kleber and Johnson(2010)and are summarized below:2.4.1.Does the solubility of SOM,as determined by chemical (alkaline/acid)extractions,sufficiently characterize recalcitrant SOM?Traditional chemical extraction procedures leave unextractable residues that are assumed to be resistant or recalcitrant because of their complex polymeric macromolecular structure. This observation led to the development of the‘humus con-cept’which postulates that decomposition processes create humic substances having different turnover times,i.e.fulvic acids(decades/centuries),humins and humic acids(millennia) (Schlesinger,1977).However,such inherent chemical stability can be questioned since at least some extracted humic substances are a product of the extraction procedure rather than a real component of SOM(Piccolo,2002).Quickly developing technologies such as nuclear magnetic resonance(NMR)and synchrotron-based,near-edge X-rayfine structure spectroscopy do not show clear evidence of discrete humic molecules in undisturbed soil(Lehmann et al., 2008b).Applying Curie point pyrolysis-gas chromatography cou-pled on-line to mass spectrometry(Py-GC/MS)and isotope ratio mass spectrometry(Py-GC IRMS),Gleixner et al.(1999)showed that instead of macromolecules,substances with lower molecular weight(proteins or peptides)are more likely to be preserved in soil during decomposition and humification processes.In addition, the assumed existence of a high proportion of aromatic C in humic materials has been questioned(Gleixner et al.,2002).These newfindings support microbial activity as the primary active agent for SOM stabilization,and that C integrated into new, microbe-derived molecules is what remains in the soil and not the precursor substances(see Chabbi and Rumpel,2009).2.4.2.Is distinction of SOM in terms of soil function and as conceptualized in process models of SOM dynamics,driven by knowledge of rates of decomposition and humification pathways?Non-living SOM is now conventionally divided into at least three C pools:(1)an active pool with turnover rates of years(root exu-dates,rapidly decomposed components of fresh plant litter)(2)an intermediate or slow pool with turnover rates of decades and(3) a passive pool with turnover rates of centuries to millennia(sta-bilized organic matter due to chemical or physical mechanisms) (Trumbore,1997,2009),including charcoal formed through pyrol-ysis.Excluding charcoal,evidence for the nature of the passive pool relies mostly on radiocarbon dating and detection of C3-plant residues in SOM of soils that have been cultivated with C4-plants for centuries or longer.The magnitude of this passive or inert pool may be grossly underestimated,if charcoal is not quantified separately (Lehmann et al.,2008a).Counter-evidence for the existence of a passive pool is that major organic materials(e.g.lignin,cellulose and hemicellulose,。

森林生态学碳固定关键科学问题

森林生态学碳固定关键科学问题

森林生态学碳固定关键科学问题英文回答:Key scientific questions in forest ecology carbon sequestration:1. How does forest structure and composition influence carbon sequestration? Forests are composed of various tree species with different growth rates and carbon storage capacities. Understanding how these factors interact and influence carbon sequestration is crucial. For example, a mixed-species forest with a diverse range of tree sizes and ages may have higher carbon sequestration rates compared to a monoculture forest.2. What are the effects of disturbance events on carbon sequestration in forests? Forests are subject to various disturbances such as wildfires, insect outbreaks, and logging activities. These disturbances can release large amounts of carbon stored in vegetation and soils. Studyingthe recovery process and the long-term effects of disturbances on carbon sequestration can provide insights into forest management strategies. For instance, a study on the impact of wildfire on carbon sequestration in a pine forest may reveal the importance of post-fire reforestation efforts.3. How do climate change and atmospheric CO2 concentrations affect forest carbon sequestration? Rising atmospheric CO2 levels and changing climatic conditions can influence forest productivity and carbon sequestration. Increased CO2 concentrations may enhance photosynthesis and promote tree growth, leading to higher carbon sequestration rates. However, the effects of climate change on forest carbon dynamics can be complex and vary across different forest types and regions. For example, a study on the response of a tropical rainforest to changing precipitation patterns and CO2 levels may reveal the potential impacts on carbon sequestration in these ecosystems.4. What is the role of soil carbon in forest carbon sequestration? Soils play a crucial role in carbonsequestration, as they can store large amounts of organic carbon. Understanding the processes that control soil carbon storage and turnover in forests is essential. For instance, a study on the effects of land management practices, such as forest thinning or organic matter additions, on soil carbon sequestration can provideinsights into sustainable forest management strategies.5. How do forest disturbances and management practices interact with carbon sequestration? Forest management practices, such as timber harvesting or afforestation, can influence carbon sequestration rates. Additionally, disturbances like fire or insect outbreaks can affect the success of management interventions. Understanding the interactions between forest disturbances and management practices is crucial for optimizing carbon sequestration efforts. For example, a study on the combined effects of timber harvesting and wildfire on carbon sequestration in a managed forest can inform sustainable forest management strategies.中文回答:森林生态学碳固定的关键科学问题:1. 森林结构和组成如何影响碳固定?森林由不同生长速度和碳储存能力的树种组成。

海獭对世界的影响英语作文

海獭对世界的影响英语作文

Sea otters, though small in stature, have a significant impact on the world in various ways. As keystone species, they play a crucial role in maintaining the balance of their ecosystems. Heres how these charming marine mammals influence the world around them:1. Ecosystem Engineers: Sea otters are known as ecosystem engineers due to their ability to shape their environment. By preying on sea urchins, they prevent these creatures from overgrazing kelp forests. Healthy kelp forests are vital as they provide habitat and food for numerous marine species, thus supporting biodiversity.2. Biodiversity Enhancement: By controlling the population of sea urchins, sea otters indirectly protect a variety of marine life that depends on kelp forests. This includes fish, invertebrates, and other organisms that call these underwater forests home.3. Carbon Sequestration: Kelp forests are efficient at capturing and storing carbon dioxide from the atmosphere. By preserving these ecosystems, sea otters contribute to the mitigation of climate change through the process of carbon sequestration.4. Economic Impact: The presence of sea otters can have economic benefits for coastal communities. Healthy marine ecosystems support fisheries and tourism, both of which can be negatively impacted by the loss of kelp forests.5. Cultural Significance: Sea otters hold cultural importance for many indigenous communities. They are often featured in traditional stories, rituals, and crafts, contributing to the rich cultural heritage of these communities.6. Conservation Efforts: The protection of sea otters has led to increased awareness and conservation efforts for marine life in general. Their status as an endangered species has prompted research and protective measures that benefit the broader marine ecosystem.7. Educational Value: Sea otters are popular animals in educational settings. They serve as ambassadors for marine conservation, teaching people about the interconnectedness of ecosystems and the importance of protecting our oceans.8. Scientific Research: Studies on sea otters have contributed to our understanding of marine biology, predatorprey dynamics, and the effects of human activity on wildlife populations.In conclusion, sea otters are more than just adorable creatures they are vital contributors to the health and balance of marine ecosystems. Their presence or absence can havefarreaching effects on the biodiversity, climate, economy, and culture of our world. As stewards of the planet, it is our responsibility to protect and preserve these remarkable animals and their habitats.。

Soil Biodiversity and Ecosystem Function

Soil Biodiversity and Ecosystem Function

Soil Biodiversity and Ecosystem Function Soil biodiversity plays a crucial role in maintaining ecosystem function and overall environmental health. The diversity of organisms living in the soil, including bacteria, fungi, protozoa, nematodes, and earthworms, contributes to various ecosystem processes such as nutrient cycling, soil structure formation, and the regulation of plant growth. However, soil biodiversity is increasingly threatened by human activities such as deforestation, intensive agriculture, urbanization, and pollution. These threats have the potential to disrupt soil ecosystems and have far-reaching consequences for the health of the planet. Oneof the key functions of soil biodiversity is nutrient cycling. Soil organisms play a vital role in breaking down organic matter and releasing nutrients such as nitrogen, phosphorus, and potassium, which are essential for plant growth. For example, mycorrhizal fungi form symbiotic relationships with plant roots, facilitating the uptake of nutrients from the soil. Additionally, earthworms and other soil-dwelling organisms help to mix and aerate the soil, improving its structure and enhancing its ability to retain water and support plant growth. Without a diverse community of soil organisms, these essential processes would be disrupted, leading to decreased soil fertility and productivity. In addition to nutrient cycling, soil biodiversity also contributes to the regulation of greenhouse gas emissions and the sequestration of carbon. Soil microorganisms, such as bacteria and fungi, play a crucial role in the decomposition of organic matter and the release of carbon dioxide, methane, and nitrous oxide. At the same time, soil can act as a carbon sink, with organic matter being stored in the soil for long periods of time. The balance between these processes is delicate and relies on the presence of a diverse array of soil organisms. Disturbances to soil biodiversity can upset this balance, potentially leading to increased greenhouse gas emissions and contributing to climate change. Furthermore, soil biodiversity is essential for supporting above-ground biodiversity. Many plant species rely on interactions with soil organisms for their growth and survival. For example, certain plant species have specific relationships with mycorrhizal fungi, which help them access nutrients from the soil. In turn, the diversity of plant species supports a wide range of above-ground organisms, including insects, birds, andmammals. Therefore, disruptions to soil biodiversity can have cascading effects on the entire ecosystem, leading to declines in above-ground biodiversity and the services it provides. Human activities have had a significant impact on soil biodiversity, with widespread consequences for ecosystem function. Deforestation, for example, not only removes above-ground vegetation but also disrupts the complex networks of soil organisms that rely on these plants for food and habitat. Similarly, intensive agricultural practices, such as the heavy use of chemical fertilizers and pesticides, can harm soil organisms and reduce overall biodiversity. Urbanization and land degradation further contribute to the loss of soil biodiversity, as natural habitats are fragmented and destroyed. In order to address these threats to soil biodiversity and ecosystem function, it is crucial to implement sustainable land management practices. This includes reducing deforestation and promoting reforestation efforts, adopting agroecological approaches to agriculture that minimize the use of chemicals and preserve natural habitats, and implementing measures to prevent soil erosion and degradation in urban areas. Additionally, raising awareness about the importance of soil biodiversity and its role in supporting ecosystem function is essential for garnering support for conservation efforts. In conclusion, soil biodiversity is a fundamental component of ecosystem function, playing a critical role in nutrient cycling, carbon sequestration, and the support of above-ground biodiversity. However, human activities pose significant threats to soil biodiversity, with far-reaching consequences for the health of the planet. By implementing sustainable land management practices and raising awareness about the importance of soil biodiversity, we can work towards preserving this vital resource and ensuring the continued health and functioning of ecosystems.。

Carbon sequestration

Carbon sequestration

, published 27 February 2008, doi: 10.1098/rstb.2007.2185363 2008 Phil. Trans. R. Soc. BRattan LalCarbon sequestrationReferences/content/363/1492/815.full.html#related-urlsArticle cited in:/content/363/1492/815.full.html#ref-list-1This article cites 115 articles, 37 of which can be accessed free Email alerting servicehere right-hand corner of the article or click Receive free email alerts when new articles cite this article - sign up in the box at the top/subscriptions go to: Phil. Trans. R. Soc. B To subscribe toCarbon sequestrationRattan Lal *Carbon Management and Sequestration Center,The Ohio State University,Columbus,OH 43210,USA Developing technologies to reduce the rate of increase of atmospheric concentration of carbon dioxide (CO 2)from annual emissions of 8.6Pg C yr –1from energy,process industry,land-use conversion and soil cultivation is an important issue of the twenty-first century.Of the three options of reducing the global energy use,developing low or no-carbon fuel and sequestering emissions,this manuscript describes processes for carbon (CO 2)sequestration and discusses abiotic and biotic technologies.Carbon sequestration implies transfer of atmospheric CO 2into other long-lived global pools including oceanic,pedologic,biotic and geological strata to reduce the net rate of increase in atmospheric CO 2.Engineering techniques of CO 2injection in deep ocean,geological strata,old coal mines and oil wells,and saline aquifers along with mineral carbonation of CO 2constitute abiotic techniques.These techniques have a large potential of thousands of Pg,are expensive,have leakage risks and may be available for routine use by 2025and beyond.In comparison,biotic techniques are natural and cost-effective processes,have numerous ancillary benefits,are immediately applicable but have finite sink capacity.Biotic and abiotic C sequestration options have specific nitches,are complementary,and have potential to mitigate the climate change risks.Keywords:climate change;greenhouse effect;soil management;geological sequestration;chemical sequestration;oceanic sequestration1.INTRODUCTIONGlobal surface temperatures have increased by 0.88C since the late nineteenth century,and 11out of the 12warmest years on record have occurred since 1995(IPCC 2007).Earth’s mean temperature is projected to increase by 1.5–5.88C during the twenty-first century (IPCC 2001).The rate of increase in global temperature has been 0.158C per decade since 1975.In addition to the sea-level rise of 15–23cm during the twentieth century (IPCC 2007),there have been notable shifts in ecosystems (Greene &Pershing 2007)and frequency and intensity of occurrence of wild fires (Running 2006;Westerling et al.2006).These and other observed climate changes are reportedly caused by emission of greenhouse gases (GHGs)through anthropogenic activities including land-use change,deforestation,biomass burning,draining of wetlands,soil cultivation and fossil fuel combustion.Consequently,the concentration of atmospheric GHGs and their radiative forcing have progressively increased with increase in human population,but especially so since the onset of industrial revolution around 1850.The concen-tration of carbon dioxide (CO 2)has increased by 31%from 280ppmv in 1850to 380ppmv in 2005,and is presently increasing at 1.7ppmv yr K 1or 0.46%yr K 1(WMO 2006;IPCC 2007).Concen-trations of methane (CH 4)and nitrous oxide (N 2O)have also increased steadily over the same period (IPCC 2001,2007;Prather et al.2001;WMO 2006).Total radiative forcing,externally imposed pertur-bation in the radiative energy budget of the Earth’sclimate system (Ramaswamy et al.2001),of all GHGs since 1850is estimated at 2.43W m K 2(IPCC 2001,2007).There is a strong interest in stabilizing the atmos-pheric abundance of CO 2and other GHGs to mitigate the risks of global warming (Kerr 2007;Kintisch 2007b ;Kluger 2007;Walsh 2007).There are three strategies of lowering CO 2emissions to mitigate climate change (Schrag 2007):(i)reducing the global energy use,(ii)developing low or no-carbon fuel,and (iii)sequestering CO 2from point sources or atmosphere through natural and engineering techniques.Between 1850and 1998,anthropogenic emissions are estimated at 270G 30Pg by fossil fuel combustion and at 136G 30Pg by land-use change,deforestation and soil cultivation (IPCC 2001).Presently,approximately 7Pg C yr K 1is emitted by fossil fuel combustion (Pacala &Socolow 2004)and 1.6Pg C yr K 1by deforestation,land-use change and soil cultivation.Of the total anthropogenic emissions of 8.6Pg C yr K 1,3.5Pg C yr K 1is absorbed by the atmosphere,2.3Pg C yr K 1by the ocean and the remainder by an unidentified terrestrial sink probably in the Northern Hemisphere (T ans et al.1990;Fan et al.1998).The objective of this paper is to discuss the process and technological options of CO 2–C sequestration in one of the long-lived global C pools so as to reduce the net rate of increase of atmospheric concentration of CO 2.While CO 2–C sequestration is discussed in general,specific attention is given to the terrestrial C sequestration in forests and soils.2.THE GLOBAL CARBON CYCLEThe importance of atmospheric concentration of CO 2on global temperature was recognized byArrhenius Phil.Trans.R.Soc.B (2008)363,815–830doi:10.1098/rstb.2007.2185Published online 30August 2007One contribution of 15to a Theme Issue ‘Sustainable agriculture II’.*lal.1@815This journal is q 2007The Royal Society(1896)towards the end of the nineteenth century, whereas anthropogenic perturbation of the global C cycle during the twentieth century has been an historically unprecedented phenomenon.Understand-ing the global C cycle and its perturbation by anthropogenic activities is important for developing viable strategies for mitigating climate change.The rate of future increase in atmospheric CO2concen-tration will depend on the anthropogenic activities, the interaction of biogeochemical and climate pro-cesses on the global C cycle and interaction among principal C pools.There arefive global C pools,of which the largest oceanic pool is estimated at 38000Pg and is increasing at the rate of 2.3Pg C yr K1(figure1).The geological C pool, comprising fossil fuels,is estimated at4130Pg,of which85%is coal, 5.5%is oil and 3.3%is gas. Proven reserves of fossil fuel include678Pg of coal (3.2Pg yr K1production),146Pg of oil(3.6Pg yr K1 of production)and98Pg of natural gas(1.5Pg yr K1 of production;Schrag2007).Presently,coal and oil each account for approximately40%of global CO2 emissions(Schrag2007).Thus,the geological pool is depleting,through fossil fuel combustion,at the rate of7.0Pg C yr K1.The third largest pool is pedologic, estimated at2500Pg to1m depth.It consists of two distinct components:soil organic carbon(SOC)pool estimated at1550Pg and soil inorganic carbon(SIC) pool at950Pg(Batjes1996).The SOC pool includes highly active humus and relatively inert charcoal C.It comprises a mixture of:(i)plant and animal residues at various stages of decomposition;(ii)substances synthesized microbiologically and/or chemically from the breakdown products;and(iii)the bodies of live micro-organisms and small animals and their decom-posing products(Schnitzer1991).The SIC pool includes elemental C and carbonate minerals such as calcite,dolomite and gypsum,and comprises primary and secondary carbonates.The primary carbonates are derived from the weathering of parent material.In contrast,the secondary carbonates are formed through the reaction of atmospheric CO2with Ca C2 and Mg C2brought in from outside the local ecosystem(e.g.calcareous dust,irrigation water, fertilizers,manures).The SIC is an important constituent in soils of arid and semi-arid regions. The fourth largest pool is the atmospheric pool comprising760Pg of CO2–C,and increasing at the rate of3.5Pg C yr K1or0.46%yr K1.The smallest among the global C pools is the biotic pool estimated at560Pg.The pedologic and biotic C pools together are called the terrestrial C pool estimated at approximately2860Pg.The atmospheric pool is also connected to the oceanic pool which absorbs 92.3Pg yr K1and releases90Pg yr K1with a net positive balance of2.3Pg C yr K1.The oceanic pool will absorb approximately5Pg C K1yr K1by2100 (Orr et al.2001).The total dissolved inorganic C in the oceans is approximately59times that of the atmosphere.On the scales of millennia,the oceans determine the atmospheric CO2concentration,not vice versa(Falkowski et al.2000).The link between geological(fossil fuel)and atmospheric pool is in one direction:transfer of approximately7.0Pg C yr K1Figure1.Principal global C pools andfluxes between them.The data on C pools among major reservoirs are from Batjes(1996), Falkowski et al.(2000)and Pacala&Socolow(2004),and the data onfluxes are from IPCC(2001).l Carbon sequestrationPhil.Trans.R.Soc.B(2008)from fossil fuel consumption to the atmosphere. The rate of fossil fuel consumption may peak by about2025.The terrestrial and atmospheric C pools are strongly interacting with one another(figure2).The annual rate of photosynthesis is120Pg C,most of which is returned back to the atmosphere through plant and soil respiration.The terrestrial C pool is depleted by conversion from natural to managed ecosystems, extractive farming practices based on low external input and soil degrading land use.The pedologic pool loses0.4–0.8Pg C yr K1to the ocean through erosion-induced transportation to aquatic ecosystems.The terrestrial sink is presently increasing at a net rate of 1.4G0.7Pg C yr K1.Thus,the terrestrial sink absorbs approximately2–4Pg C yr K1and its capacity may increase to approximately5Pg C yr K1by2050 (Cramer et al.2001;Scholes&Noble2001).Increase in the terrestrial sink capacity may be due to CO2 fertilization effect and change in land use and manage-ment.The biotic pool also contributes to increase in atmospheric CO2concentration through deforestation and biomass burning.3.CARBON SEQUESTRATIONEmission rates from fossil fuel combustion increased by40%between1980and2000(Wofsy2001).Y et, the amount of CO2accumulating in the atmosphere remained the same over this period because the excess CO2released is being removed by oceans,forests,soils and other ecosystems(Battle et al.2000).Atmospheric CO2increased at a rate of2.8–3.0Pg C yr K1during the1980s and1995,and between 3.0and 3.5Pg C yr K1during1995–2005.Considering the total anthropogenic emissions of between6and 8Pg C yr K1,while the atmospheric increase has beenFigure2.The atmospheric C pool is increasing at the rate of3.5Pg C yr K1.The terrestrial C pool contributes approximately 1.6Pg C yr K1through deforestation,biomass burning,draining of wetlands,soil cultivation including those of organic soils, accelerated erosion and hidden C costs of input(e.g.fertilizers,tillage,pesticides,irrigation).T errestrial C pools are presently sink of2–4Pg C yr K1.Conversion to a judicious land use and adoption of recommended practices in managed ecosystems can make these important sinks especially due to CO2fertilization effects.Phil.Trans.R.Soc.B(2008)2.8–3.5Pg yr K1,implies the existence of large global terrestrial sinks(Fung2000;Pacala2001).Fan et al. (1998)estimated a mean annual uptake of 1.7G 0.5Pg C yr K1in North America,mostly south of 518N.In comparison,the Eurasia–North Africa sink was relatively small.This process of transfer and secure storage of atmospheric CO2into other long-lived C pools that would otherwise be emitted or remain in the atmosphere is called‘carbon sequestra-tion’.Therefore,in this context,C sequestration may be a natural or an anthropogenically driven process. The objective of an anthropogenically driven C sequestration process is to balance the global C budget such that future economic growth is based on a‘C-neutral’strategy of no net gain in atmospheric C pool. Such a strategy would necessitate sequestering almost all anthropogenically generated CO2through safe, environmentally acceptable and stable techniques with low risks of ckner(2003)estimated that if a C-neutral strategy is based mainly on sequestration rather than emission reduction,total C storage during the twenty-first century will exceed600Pg with residence time of centuries to millennia.However, even a small leakage of2–3Pg C yr K1from the C sequestered in one of the pools can adversely impact the strategic planning for future generations.Pacala& Socolow(2004)outlined15options of stabilizing the atmospheric concentration of CO2by2050at approximately550ppm.Of the15options,three were based on C sequestration in terrestrial ecosystems.There are several technological options for seques-tration of atmospheric CO2into one of the other global pools(figure3).The choice of one or a combination of several technologies is important for formulating energy policies for future economic growth and development at national and global scales.These options can be grouped into two broad categories: abiotic and biotic sequestration.(a)Abiotic sequestrationAbiotic sequestration is based on physical and chemical reactions and engineering techniques without inter-vention of living organisms(e.g.plants,microbes).The abiotic strategy of C sequestration in oceanic and geological structures has received considerable atten-tion(Freund&Ormerod1997)because theoretically abiotic sequestration has a larger sink capacity than biotic sequestration.Rapid progress is being made in developing/testing technologies for CO2capture, transport and injection(Kerr2001).(i)Oceanic injectionInjection of a pure CO2stream deep in the ocean is a possibility that has been widely considered by engineers for about three decades.Following the initial proposal in the late1970s,there has been considerable progress in oceanic injection of CO2.T o be stable and minimize outgassing,CO2must be injected at great depths. Therefore,liquefied CO2separated from industrial sources can be injected into the ocean by one of the following four techniques:(i)it is injected below 1000m from a manifold lying at the oceanfloor,and being lighter than water,it rises to approximately 1000m depth forming a droplet plume;(ii)it is also injected as a denser CO2–seawater mixture at 500–1000m depth,and the mixture sinks into the deeper ocean;(iii)it is discharged from a large pipe towed behind a ship;and(iv)it is pumped into a depression at the bottom of the oceanfloor forming a CO2lake.Liquefied CO2injected at approximately 3000m depth is believed to remain stable(O’Connor et al.2001).The oceanic sink capacity for CO2 sequestration is estimated at5000–10000Pg C,Figure3.A wide range of processes and technological options for C sequestration in agricultural,industrial and natural ecosystems.l Carbon sequestrationPhil.Trans.R.Soc.B(2008)exceeding the estimated fossil fuel reserves(Herzog et al.1997,2002).However,CO2injection may also have some adverse effects on deep sea biota(Auerbach et al.1997;Caulfield et al.1997;Seibel&Walsh2001). In addition to the economics,the issue of stability of such an injection must be addressed owing to the increased stratification of the ocean water column and its turn over through natural processes.(ii)Geological injectionThis involves capture,liquefaction,transport and injection of industrial CO2into deep geological strata. The CO2may be injected in coal seams,old oil wells(to increase yield),stable rock strata or saline aquifers (Tsang et al.2002;Klara et al.2003;Baines&Worden 2004;Gale2004).Saline aquifers are underground strata of very porous sedimentsfilled with brackish (saline)water.In general,saline aquifers are located below the freshwater reservoirs with an impermeable layer in between.Industrial CO2can be pumped into the aquifer,where it is sequestered hydrodynamically and by reacting with other dissolved salts to form carbonates.Carbon dioxide is injected in a supercritical state that has much lower density and viscosity than the liquid brine that it displaces.In situ,it forms a gas-like phase and also dissolves in the aqueous phase,creating a multiphase multicomponent environment.Injecting CO2into reservoirs in which it displaces oil or gas could be an economic strategy of enhanced oil recovery (EOR).Production from oil and gasfields,which has been in decline,is raised by CO2-enhanced recovery (Klusman2003).This strategy of CO2sequestration is used in T exas,USA,to inject20million Mg of CO2yr K1at a price of$10–$15Mg K1(Lackner 2003).The technique is also used in offshore oil wells in Norway.In a strict sense,however,this is not sequestration when the CO2injected is extracted from underground wells.The CO2can also be injected into unmineable coal seams where CH4is absorbed. Injected CO2is absorbed onto coal twice as much as CH4,and the process enhances the gas recovery of coal bed CH4(CBM).Principal concerns about geological sequestration,similar to that of the oceanic,are (Kintisch2007a;Schrag2007):(i)reliability of storage of vast quantities of CO2in geological strata and(ii)the cost.Some have argued that risks of leakage are low. However,to date,a few direct injection of CO2have been made on a commercial scale although the Regional Carbon Sequestration Partnership project of US DOE is planning for several demonstrations during 2008–2009.Similar to oceanic injection,cost and leakage are principal issues of geological sequestration which need to be resolved.Owing to the low density and viscosity and injection under supercritical con-ditions,the risks of CO2leakage through confining strata may be higher than currently injected liquid wastes(Tsang et al.2002).In addition,the chemical interactions of CO2with the geological formations may have to be considered in formulating guidelines for appropriate regulatory and monitoring controls. (iii)Scrubbing and mineral carbonationMineral carbonation is achieved through mimicry of natural inorganic chemical transformation of CO2(Fan&Park2004).It involves transformation of industrial strength CO2emissions into CaCO3, MgCO3and other minerals in the form of geologically and thermodynamically stable mineral carbonates.It is a two-stage process:scrubbing and mineral carbona-tion.Scrubbing,the process of chemical absorption of CO2using an amine or carbonate solvent,is the most widely used method of carbon capture.The CO2is purified by passing through an absorption column containing amine solvent.Other solvents used are K2CO3,lithium silicate,ceramic and nickel-based compounds.Pure CO2gas,recovered by heating the CO2-rich amine,is re-precipitated through mineral carbonation.Carbonates thus formed are stable rock in which CO2is sequestered forever.Aqueous mineral carbonation reactions leading to the formation of magnesite(MgCO3),olivine(Mg2SiO4)and serpen-tine(Mg3Si2O5(OH)4)are as follows(Gerdemann et al.2003):2MgSiO4C CO2ðgÞC2H2O/Mg3Si2O5ðOHÞ4C MgCO3ð16:5KCalÞ;Mg2SiO4C2CO2ðgÞ/2MgCO3C SiO2ð10:3KCalÞandCaSiO3C CO2/CaCO3C SiO2ð10:6KCalÞ:All these reactions occur in nature and can be replicated in industrial settings.The industrial process of mineral carbonation has been described by Lackner et al.(1996,1997)and O’Connor et al.(2000).The process uses a slurry offine particle-sized mineral in water,at solid concentrations of15–30%.Dissolution of the mineral and subsequent carbonation occur in a single operation as per the following theorized reactions (O’Connor et al.2000;Gerdemann et al.2004): CO2C H2O/H2CO3/H C C HCO K3;Mg2SiO4C4H C/2Mg C2C H4SiO4or SiO2C2H2O andMg C2C HCO K3/MgCO3C H2:Geological studies have shown that reserves of ultramafic(ultrabasic)minerals are sufficient to provide raw materials for mineral carbonation of industrial emissions for long time(Goff et al.1997, 2000).However,these are geological reactions and occur at slow rate.The challenge lies in increasing the rate of reaction by decreasing the particle size, increasing temperature and pressure and using catalytic agents(Fan&Park2004).Increasing the rate of reactions,however,requires energy and is expensive.(b)Biotic sequestrationBiotic sequestration is based on managed intervention of higher plants and micro-organisms in removing CO2 from the atmosphere.It differs from management options which reduce emission or offset emission. Increasing use efficiency of resources(e.g.water, energy)is another option for managing the terrestrial C pool(table1).Some biotic sequestration options are briefly described below.Carbon sequestration l819Phil.Trans.R.Soc.B(2008)(i)Oceanic sequestrationThere are several biological processes leading to C sequestration in the ocean through photosynthesis. Phytoplankton photosynthesis is one such mechanism (Rivkin&Legendre2001),whichfixes approximately 45Pg C yr K1(Falkowski et al.2000).Some of the particulate organic material formed by phytoplankton is deposited at the oceanfloor and is thus sequestered (Raven&Falkowski1999).Availability of Fe is one of the limiting factors on phytoplankton growth in oceanic ecosystems.Thus,several studies have assessed the importance of Fe fertilization on biotic CO2sequestra-tion in the ocean(Martin&Fitzwater1988;Falkowski 1997;Martin et al.2002;Boyd et al.2004).It is also argued that incremental C could be sold as credits in the developing global C marketplace.Similar to deep injection,ocean fertilization may also change the ecology of the ocean(Chisholm et al.2001).However, with the current state of knowledge,the topic of ocean fertilization remains a debatable issue(Johnson& Karl2002).(ii)T errestrial sequestrationTransfer of atmospheric CO2into biotic and pedologicC pools is called terrestrial C sequestration.Of the8.6Pg C yr K1emitted into the atmosphere,only 3.5Pg or40%of the anthropogenically emitted CO2 remains in the atmosphere primarily owing to unspe-cified terrestrial sinks which sequester atmospheric CO2and play an important role in the global C cycle. T errestrial ecosystems constitute a major C sink owing to the photosynthesis and storage of CO2in live and dead organic matter.Owing to its numerous ancillary benefits(e.g.improved soil and water quality,restor-ation of degraded ecosystems,increased crop yield), terrestrial C sequestration is often termed as win–win or no-regrets strategy(Lal et al.2003).It offers multiple benefits even without the threat of global climate change.There are three principal components of terrestrial C sequestration:forests, soils and wetlands.Forest ecosystems store C as lignin and other relatively resistant polymeric C compounds.Presently,the net rate of C sequestration in forest ecosystems(other than those being deforested)is 1.7G0.5Pg C yr K1(Fan et al.1998).The forest C is sequestered not only in the harvestable timber,but also in woody debris, wood products and other woody plants encroaching upon grasslands(Wofsy2001).The terrestrial NPP is not saturated at the current CO2concentration and may increase with increase in atmospheric CO2 concentration through the CO2fertilization effect. The NPP may be saturated at800–1000ppm of CO2 concentration(Falkowski et al.2000).Thus,the forest sink may increase by CO2fertilization effect (Krishnamurthy&Machavaram2000).The potential CO2fertilization effect may peak during the middle of the twenty-first century(Kohlmaier et al.1998). However,at increasing CO2concentration,the NPP may be limited by the deficiency of N,P,H2O and other factors.Interaction between cycles of C,N,P and H2O, if moderated through judicious management,may enhance terrestrial C sequestration.Afforestation is one of the viable options of C sequestration in terrestrial ecosystems(IPCC1999; Fang et al.2001;Lamb et al.2005).The potential of C sequestration through afforestation is estimated at 3Tg C yr K1in Norway,6Tg C yr K1in New Zealand, 9Tg C yr K1in Sweden,107Tg C yr K1in Russia and 117Tg C yr K1in the USA(IPCC1999).The rate of C sequestration in US forests,considering all com-ponents,is0.3–0.7Pg C yr K1(Pacala et al.2001). Restoration of degraded tropical forest is another important option(Lamb et al.2005).It is estimated that350Mha of tropical forest have been converted to other land uses,and another500Mha of forests have been degraded to varying extent(Lal2005a,b,c).Thus, establishment of productive and monoculture planta-tions of Pinus,Eucalyptus and Acacia can enhance the terrestrial C pool in these ecosystems.There is also a potential for improving the management of secondary or regrowth forests in degraded tropical landscapes. Fang et al.(2001)estimated that in China,between 1970and1998,C sequestration in forests increased at an average rate of21Tg C yr K1mainly due to afforestation and growth.The total C pool in Chinese forest decreased from5.1Pg in1949to4.3Pg in1977–1981.Subsequently,it progressively increased during 1980s and1990s to4.7Pg in1998(Fang et al.2001). T ownsend et al.(2002)suggested a sizeable CO2T able1.T errestrial carbon management options.management of terrestrial C pool sequestration of C in terrestrial pool reducing emissions sequestering emissions as SOC eliminating ploughing increasing humification efficiency conserving water and decreasing irrigation need improving soil aggregationusing integrated pest management to minimize the use of pesticides deep incorporation of SOC through establishing deep-rooted plants,promoting bioturbation and transfer of DOC into the ground waterbiological nitrogenfixation to reduce fertilizer useoffsetting emissions sequestering emissions as SICestablishing biofuel plantations forming secondary carbonates through biogenic processes biodigestion to produce CH4gas leaching of biocarbonates into the ground waterbio-diesel and bioethanol productionenhancing use efficiencyprecision farmingfertilizer placement and formulationsdrip,sub-irrigation or furrow irrigationl Carbon sequestrationPhil.Trans.R.Soc.B(2008)uptake in C-3dominated tropical regions in eight of the 10-year study period.Their data showed a possible existence of a large equatorial terrestrial CO2sink.Pacala&Socolow’s(2004)estimated management of temperate and tropical forest is one of the15options to stabilize atmospheric CO2concentration at550ppm by2050.They estimated that0.5Pg C emission would be avoided if the current rate of clear cutting of primary tropical forest were reduced to zero over50years by 2050.Another0.5Pg C yr K1would be sequestered by reforesting or afforesting approximately250Mha in the tropics or400Mha in the temperate zone.An additional0.3Pg C yr K1can be sequestered by estab-lishing approximately300Mha of plantations of non-forested lands(Pacala&Socolow2004).The afforestation may account for a total of25Pg C sequestration between2000and2050.However,afforestation on a large scale can impact water resources.Jackson et al.(2005)documented substantial losses in streamflow,and increased soil salinization and acidification,with afforestation.They observed that establishment of tree plantations decreased streamflow by227mm yr K1globally (52%),with13%of streams drying completely for at least one year.Thus,any plans of large-scale afforesta-tion for C sequestration must consider the possible adverse impact on availability of water.Bunker et al.(2005)addressed the concern of decline in tropical forest biodiversity due to expansion of monoculture plantations.Thus,resulting effects of monoculture plantations on key ecosystem services (e.g.biodiversity,water resources,elemental cycling,C sequestration)must be critically assessed.There is a strong need for developing regulatory policies,includ-ing the transaction costs of trading C credits and obtaining the permits.The cost of C sequestration vis-a`-vis the opportunity cost must also be considered (McCarl&Schneider2001).Wetlands and the associated soils or histosols constitute a large pedologic pool estimated at approxi-mately450Pg(Gorham1991;Warner et al.1993). Wetland soils may contain as much as200times more C than the associated vegetation(Milne&Brown 1997;Garnett et al.2001).Gorham(1991)and Kobak et al.(1998)estimated that C sequestration in wet-lands/peatsoils since the post-glaciation period resulted in the C accumulation at the rate of0.1Pg C yr K1over 10000–18000years.However,drainage of peatlands and their subsequent cultivation made these ecosys-tems a net source of rge areas of wetlands have been drained worldwide for agriculture(Armentano& Menges1986)and forestry(Paavilainen&Paivanen 1995).Drained wetland soils decompose and subside at the rate of approximately1–2cm yr K1primarily due to oxidation(Rojstaczer&Deverel1995;Hillman 1997;Wosten et al.1997).Restoration of wetlands can lead to reversal of the process and make restored wetlands once again a sink of atmospheric CO2. However,there is a long time lag after the restoration until processes in restored wetlands become similar to those of natural wetlands.Soil C sequestration.Implies enhancing the concen-tration/pools of SOC and SIC as secondary carbo-nates through land-use conversion and adoption of recommended management practices(RMPs)in agricultural,pastoral and forestry ecosystems and restoration of degraded and drastically disturbed soils.Formation of charcoal and use of biochar as a fertilizer is another option(Fowles2007).In contrast to geological sequestration that implies injecting CO2 at1–2km depth,the SOC sequestration involves putting C into the surface layer of0.5–1m depth through the natural processes of humification.Most soils under the managed ecosystems contain a lower SOC pool than their counterparts under natural ecosystems owing to the depletion of the SOC pool in cultivated soils.The most rapid loss of the SOC pool occurs in thefirst20–50years of conversion of natural to agricultural ecosystems in temperate regions and 5–10years in the tropics(Lal2001).In general, cultivated soils normally contain50–75%of the original SOC pool.The depletion of the SOC pool is caused by oxidation/mineralization,leaching and erosion.Strategies for increasing the SOC pool are outlined infigure4.There is a wide range of degraded soils with a depleted SOC pool.Important among these are those degraded by erosion,nutrient depletion,acidification and leaching,structural decline and pollution/contami-nation(figure4).Restoring degraded soils and ecosystems is a strategy with multiple benefits for water quality,biomass productivity and for reducing net CO2emission.Grainger(1995)estimated that there are approximately750Mha of degraded lands in the tropics with potential for afforestation and soil quality enhancement.With a sequestration potential of approximately0.5Mg ha K1yr K1as SOC and an additional1.0Mg ha K1yr K1as biomass,a terrestrialC sequestration potential of750Mha is approximately1.1Pg C yr l(2001)estimated the SOC seques-tration potential of0.4–0.7Pg C yr K1through deserti-fication control in soils of arid and semi-arid regions.Similar estimates were provided by Squires et al.(1995).West&Post(2002)assessed SOC sequestration rate upon conversion of plough tillage to no-till farming through analyses of data of67long-term experiments from around the world.They reported the mean rate of SOC sequestration of570G140Kg C ha K1yr K1, which may lead to the new equilibrium SOC pool in 40–l(2004a,b)estimated the global SOC sequestration potential,0.4–1.2Pg C yr K1or5–15%, of the global fossil fuel emissions.Pacala&Socolow (2004)estimated that conversion of plough tillage to no-till farming on1600Mha of cropland along with adoption of conservation-effective measures could lead to sequestration of0.5–1Pg C yr K1by2050.Integrated nutrient management(INM)is also essential to SOC sequestration.The humification process can be severely constrained by the lack of N, P,S and other building blocks of soil humus(Himes 1998).The efficiency of C sequestration is reduced when C and N are not adequately balanced(Paustian et al.1997).Therefore,the SOC sequestration rate is enhanced by an increase in the application of biomass C(Campbell et al.1991;Janzen et al.1998)and N (Halvorson et al.1999,2002).Liebig et al.(2002) observed that high N rate treatments increased SOCCarbon sequestration l821Phil.Trans.R.Soc.B(2008)。

土壤碳的固持 英语

土壤碳的固持英语Soil Carbon sequestration: An Essential Process for Sustainable Agriculture and Climate Change Mitigation.Soil carbon sequestration, also known as soil carbon storage or soil carbon fixation, is a critical process in maintaining the health and productivity of soil ecosystems. It involves the stabilization and accumulation of carbon in soil organic matter, which can occur through various biological, chemical, and physical processes. This process is crucial for sustainable agriculture, climate change mitigation, and the overall well-being of the planet.The importance of soil carbon sequestration cannot be overstated. Soils are the largest terrestrial carbon pool, storing more carbon than the atmosphere and all plant biomass combined. This carbon is stored in the form of organic matter, which is composed of dead and decaying plant and animal remains, as well as living microorganisms. When carbon is sequestered in soil organic matter, it iseffectively removed from the atmosphere, thus reducing the concentration of greenhouse gases and mitigating theimpacts of climate change.There are several key processes involved in soil carbon sequestration. One of the most important is photosynthesis, which occurs in plants and converts carbon dioxide into organic matter. When plants die and decay, their carbon-rich remains are incorporated into soil organic matter, contributing to carbon sequestration. Additionally, soil microorganisms play a crucial role in decomposing organic matter and releasing carbon dioxide, which can then be re-fixed by plants through photosynthesis.The management of agricultural systems cansignificantly impact soil carbon sequestration. For example, the use of organic matter in soil amendments, such as compost or manure, can increase the amount of carbon insoil organic matter. Conservation agriculture practices, such as no-till farming or crop rotation, can also enhance soil carbon sequestration by reducing soil erosion and increasing soil organic matter content.The benefits of soil carbon sequestration extend beyond climate change mitigation. Healthy soils are essential for sustainable agriculture, providing a foundation for crop growth and yield. Soil organic matter improves soil structure, water retention, and nutrient cycling, all of which are crucial for crop productivity. By sequestering carbon in soil organic matter, farmers can enhance the fertility and resilience of their soils, leading to improved crop yields and reduced dependence on external inputs.In addition to its agricultural benefits, soil carbon sequestration can also contribute to the achievement of global climate goals. The Intergovernmental Panel on Climate Change (IPCC) has identified soil carbon sequestration as one of the most effective and cost-efficient strategies for reducing greenhouse gas emissions and mitigating climate change. By increasing soil carbon sequestration, we can not only reduce the amount of carbon dioxide in the atmosphere but also enhance the resilience and productivity of our agricultural systems.In conclusion, soil carbon sequestration is a crucial process for sustainable agriculture, climate change mitigation, and the overall well-being of the planet. It involves the stabilization and accumulation of carbon in soil organic matter, which can be enhanced through various agricultural management practices. By sequestering carbon in soil organic matter, we can reduce greenhouse gas emissions, improve soil health and fertility, andcontribute to the achievement of global climate goals. As we face the challenges of climate change and sustainable development, soil carbon sequestration remains an essential component of our efforts to protect our planet and ensure a sustainable future for all.。

Soil Carbon Sequestration

Soil Carbon SequestrationSoil carbon sequestration is an important process that involves the capture and storage of carbon dioxide from the atmosphere into the soil. This process is vital for mitigating the effects of climate change. The ability of soil to sequester carbon depends on various factors such as soil type, land use, and management practices. In this essay, I will discuss the importance of soil carbon sequestration, the factors that affect it, and the various management practices that can enhance it.Soil carbon sequestration is an essential process that helps to reduce the amount of carbon dioxide in the atmosphere. The process involves the conversion of atmospheric carbon dioxide into organic matter, which is then stored in the soil. The carbon is stored in the soil for long periods, which helps to reduce the amount of carbon dioxide in the atmosphere. This process is critical for mitigating the effects of climate change, which is one of the most significant challenges facing humanity today.The amount of carbon that can be sequestered in the soil depends on various factors such as soil type, land use, and management practices. Different soils have different capacities to store carbon. For instance, soils with high clay content can store more carbon than sandy soils. Similarly, land use practices such as deforestation and agriculture can have a significant impact on soil carbon sequestration. Deforestation leads to the loss of trees, which are essential for carbon sequestration, while agriculture can either enhance or reduce soil carbon sequestration depending on the management practices employed.Various management practices can enhance soil carbon sequestration. One of the most effective practices is the use of cover crops. Cover crops are crops that are planted to cover the soil during fallow periods. These crops help to prevent soil erosion, increase soil organic matter, and enhance soil fertility. They also help to sequester carbon by capturing carbon dioxide from the atmosphere and converting it into organic matter, which is then stored in the soil.Another effective practice is the use of conservation tillage. Conservation tillage involves reducing the amount of tillage or eliminating it altogether. This practice helps toreduce soil disturbance, which can lead to the loss of soil organic matter and soil erosion. It also helps to increase soil water retention, which is essential for plant growth and carbon sequestration.In conclusion, soil carbon sequestration is an essential process that helps to reduce the amount of carbon dioxide in the atmosphere. The amount of carbon that can be sequestered in the soil depends on various factors such as soil type, land use, and management practices. Different management practices can enhance soil carbon sequestration, including the use of cover crops, conservation tillage, and the use of organic fertilizers. These practices are critical for mitigating the effects of climate change, which is one of the most significant challenges facing humanity today. We must adopt these practices to ensure that we leave a better planet for future generations.。

Plant-Microbe Interactions in the Rhizosphere

Plant-Microbe Interactions in theRhizospherePlant-microbe interactions in the rhizosphere play a crucial role in shaping the health and productivity of plants. The rhizosphere is the region of soil surrounding plant roots where a complex network of interactions occurs between plants, microbes, and soil particles. These interactions can be beneficial, neutral, or harmful, depending on the specific organisms involved and the environmental conditions present. Understanding the dynamics of these interactions is essential for sustainable agriculture and ecosystem management. One of the most well-known examples of beneficial plant-microbe interactions in the rhizosphere is the symbiotic relationship between plants and mycorrhizal fungi. Mycorrhizal fungi form associations with plant roots, providing the plant with increased access to nutrients such as phosphorus and nitrogen. In return, the plant supplies the fungi with sugars produced through photosynthesis. This mutualistic relationship benefits both parties and enhances the overall health and growth of the plant. In addition to mycorrhizal fungi, other beneficial microbes such as rhizobia and plant growth-promoting rhizobacteria (PGPR) play important roles in the rhizosphere. Rhizobia form symbiotic relationships with leguminous plants, fixing atmospheric nitrogen into a form that can be utilized by the plant. PGPR, on the other hand, promote plant growth through various mechanisms such as nutrient solubilization, hormone production, and disease suppression. These beneficial microbes contribute to plant health and productivity by enhancing nutrient uptake, improving stress tolerance, and protecting against pathogens. While beneficial interactions in the rhizosphere are essential for plant health, harmful interactions can also occur. Pathogenic microbes such as root rot fungi and nematodes can negatively impact plant growth and productivity by causing diseases and reducing nutrient uptake. These harmful interactions highlight the importance of maintaining a balanced microbial community in the rhizosphere to prevent the proliferation of pathogens. In addition to the direct effects on plant health, plant-microbe interactions in the rhizosphere also play asignificant role in ecosystem functioning and nutrient cycling. Microbes in therhizosphere contribute to soil organic matter decomposition, nutrient mineralization, and carbon sequestration, influencing soil fertility and overall ecosystem productivity. Understanding the intricate relationships between plants and microbes in the rhizosphere is essential for sustainable land management practices and ecosystem conservation. Overall, plant-microbe interactions in the rhizosphere are complex and dynamic, with both beneficial and harmful interactions shaping plant health, productivity, and ecosystem functioning. By studying and manipulating these interactions, researchers and farmers can enhance plant growth, improve soil health, and promote sustainable agricultural practices. Embracing the complexity of the rhizosphere and harnessing the power of beneficial microbes will be essential for meeting the challenges of food security and environmental sustainability in the face of a changing climate.。

Stabilization mechanisms of soil organic matter- Implications for C-saturation of soils

Plant and Soil241:155–176,2002.©2002Kluwer Academic Publishers.Printed in the Netherlands.155 ReviewStabilization mechanisms of soil organic matter:Implications forC-saturation of soilsJ.Six1,R.T.Conant,E.A.Paul&K.PaustianNatural Resource Ecology Laboratory,Colorado State University,Fort Collins,CO80523,U.S.A.1Corresponding author∗Received3January2001.Accepted in revised form13February2002AbstractThe relationship between soil structure and the ability of soil to stabilize soil organic matter(SOM)is a key element in soil C dynamics that has either been overlooked or treated in a cursory fashion when developing SOM models. The purpose of this paper is to review current knowledge of SOM dynamics within the framework of a newly proposed soil C saturation concept.Initially,we distinguish SOM that is protected against decomposition by various mechanisms from that which is not protected from decomposition.Methods of quantification and characteristics of three SOM pools defined as protected are discussed.Soil organic matter can be:(1)physically stabilized,or protected from decomposition,through microaggregation,or(2)intimate association with silt and clay particles, and(3)can be biochemically stabilized through the formation of recalcitrant SOM compounds.In addition to behavior of each SOM pool,we discuss implications of changes in land management on processes by which SOM compounds undergo protection and release.The characteristics and responses to changes in land use or land management are described for the light fraction(LF)and particulate organic matter(POM).We defined the LF and POM not occluded within microaggregates(53–250µm sized aggregates as unprotected.Our conclusions are illustrated in a new conceptual SOM model that differs from most SOM models in that the model state variables are measurable SOM pools.We suggest that physicochemical characteristics inherent to soils define the maximum protective capacity of these pools,which limits increases in SOM(i.e.C sequestration)with increased organic residue inputs.IntroductionMost current models of SOM dynamics assumefirst-order kinetics for the decomposition of various con-ceptual pools of organic matter(McGill,1996;Paus-tian,1994),which means that equilibrium C stocks are linearly proportional to C inputs(Paustian et al., 1997).These models predict that soil C stocks can, in theory,be increased without limit,provided that C inputs increase without limit,i.e.there are no as-sumptions of soil C saturation.While these models have been largely successful in representing SOM dynamics under current conditions and management practices(e.g.Parton et al.,1987,1994;Paustian et ∗FAX No:+1-970-491-1965.E-mail:johan@ al.,1992;Powlson et al.,1996),usually for soils with low to moderate C levels(e.g.<5%),there is some question of their validity for projecting longer term SOM dynamics under scenarios of ever increasing C inputs(e.g.Donigian et al.,1997).Such scenarios are particularly relevant with the development of new technology designed to promote soil C sequestration through increasing plant C inputs.Native soil C levels reflect the balance of C inputs and C losses under native conditions(i.e.productivity, moisture and temperature regimes),but do not neces-sarily represent an upper limit in soil C stocks.Empir-ical evidence demonstrates that C levels in intensively managed agricultural and pastoral ecosystems can ex-ceed those under native conditions.Phosphorous fer-tilization of Australian pasture soils can increase soil C by150%or more relative to the native condition156(Barrow,1969;Ridley et al.,1990;Russell1960). Soil C levels under long-term grassland(‘near native’) vegetation have also been exceeded in high productiv-ity mid-western no-tillage(NT)systems(Ismail et al., 1994)as well as in sod plots with altered vegetation (Follett et al.,1997).Hence,native soil C levels may not be an appropriate measure of the ultimate C sink capacity of soils.There are several lines of evidence that suggest the existence of a C saturation level based on physiochem-ical processes that stabilize or protect organic com-pounds in soils.While many long-termfield experi-ments exhibit a proportional relationship between C inputs and soil C content across treatments(Larson et al.,1972;Paustian et al.,1997),some experiments in high C soils show little or no increase in soil C content with two to three fold increases in C inputs (Campbell et al.,1991;Paustian et al.,1997;Solberg et al.,1997).Various physical properties(e.g.silt plus clay content and microaggregation)of soil are thought to be involved in the protection of organic materials from decomposer organism.However,these proper-ties and their exerted protection seem to be limited by their characteristics(e.g.surface area),which is con-sistent with a saturation phenomenon(Hassink,1997; Kemper and Koch,1966).A number of soil organic matter models have been developed in the last30years.Most of these mod-els represent the heterogeneity of SOM by defining several pools,typically three tofive,which vary in their intrinsic decay rates and in the factors which control decomposition rates(see reviews by McGill, 1996;Parton,et al.,1994;Paustian,1994).Alternat-ive formulations,whereby specific decomposition rate varies as a function of a continuous SOM quality spec-trum(i.e.instead of discrete pools),have also been developed(e.g.Bosatta and Agren,1996).However, in either case,the representation of the model pools (or quality spectrum)is primarily conceptual in nature. While such models can be successfully validated us-ing measurements of total organic carbon and isotopic ratios of total C(e.g.Jenkinson and Rayner,1977), the individual pools are generally only loosely associ-ated with measurable quantities obtained with existing analytical methods.Consequently,it is not straight-forward to falsify or test the internal dynamics of C transfers between pools and changes in pool sizes of the current SOM models with conceptual pool defin-itions because a direct comparison to measured pool changes is not possible.A closer linkage between theoretical and measur-able pools of SOM can be made by explicitly defining model pools to coincide with measurable quantities or by devising more functional laboratory fractiona-tion procedures or both.The phrases‘modeling the measurable’and‘measuring the modelable’have been coined as representing the two approaches towards a closer reconciliation between theoretical and experi-mental work on SOM(Christensen,1996;Elliott et al.,1996).Various attempts have been made to correlate ana-lytical laboratory fractions with conceptual model pools,with limited success.Motavalli et al.(1994) compared laboratory measurements of C mineraliza-tion with simulations by the Century model(Parton et al.,1994)for several tropical soils.When the active and slow pools in the model were initialized using laboratory determinations of microbial+soluble C for the active pool and light fraction for the slow pool,C mineralization was consistently underestim-ated,although all fractions were highly significantly correlated to C mineralization in a regression ana-lysis.Magid et al.(1996)unsuccessfully attempted to trace14C labeled plant materials using three size-density fractionation methods to define an‘active’pool.Metherell(1992)found that the slow pool in Century was much larger than the particulate organic matter(POM)fraction isolated from a Haplustoll by Cambardella and Elliott(1992).However,Balesdent (1996)found that POM isolated after mild disrup-tion corresponds to the plant structural compartment (RPM)of the Rothamsted carbon model(Jenkinson and Rayner,1997).Acid hydrolysis has been used to estimate Century’s passive C pool(Paul et al.,1997a; Trumbore,1993),but it seems to slightly overestimate the size(Paul et al.,1997a;Trumbore,1993),though not the C turnover rate,of the passive pool(Trumbore, 1993).Nevertheless,Paul et al.(1999)used extended laboratory incubations in combination with acid hy-drolysis to define an active,slow and passive pool of C and were successful in modeling the evolution of CO2in thefield based on these pools.These studies suggest that attempting to measure the modelable has had minimal success to date.There have been a few recent attempts to more closely integrate models and measurements of physi-cochemically defined pools by‘modeling the measur-able’,although Elliott et al.(1996)and Christensen (1996)have presented conceptual models for this ap-proach.Arah(2000)proposed an approach based on analytically defined pools and measurements of13C157Figure1.The protective capacity of soil(which governs the silt-and clay protected C and microaggregate protected C pools),the biochemically stabilized C pool and the unprotected C pool define a maximum C content for soils.The pool size of each fraction is determined by their unique stabilizing mechanisms.and15N stable isotope tracers to derive parameters for a model with measurable pools.The approach con-siders all possible transformations between measured C and N pools and devises a system of equations using observed changes in total C and N and13C and15N for each fraction to solve all model unknowns.Necessary requirements of such an approach are that the analyt-ical fractions are distinct and together account for the total carbon inventory.The objective of this review paper is to summar-ize current knowledge on SOM dynamics and sta-bilization and to synthesize this information into a conceptual SOM model based on physicochemically defined SOM pools.This new model defines a soil C-saturation capacity,or a maximum soil C storage potential,determined by the physicochemical proper-ties of the soil.We propose that the conceptual model developed from this knowledge may form the basis for a simulation model with physicochemically measur-able SOM pools as state variables rather than with the biologically defined pools by Paul et al.(1999).Protected SOM:Stabilization mechanisms, characteristics,and dynamicsThree main mechanisms of SOM stabilization have been proposed:(1)chemical stabilization,(2)phys-ical protection and(3)biochemical stabilization (Christensen,1996;Stevenson,1994).Chemical sta-bilization of SOM is understood to be the result of the chemical or physicochemical binding between SOM and soil minerals(i.e.clay and silt particles).Indeed, many studies have reported a relationship between stabilization of organic C and N in soils and clay or silt plus clay content(Feller and Beare,1997; Hassink,1997;Ladd et al.,1985;Merckx et al., 1985;Sorensen,1972).In addition to the clay con-tent,clay type(i.e.2:1versus1:1versus allophanic clay minerals)influences the stabilization of organic C and N(Feller and Beare,1997;Ladd et al.1992; Sorensen,1972;Torn et al.,1997).Physical protection by aggregates is indicated by the positive influence of aggregation on the accumulation of SOM(e.g.Ed-wards and Bremner,1967;Elliott,1986;Jastrow, 1996;Tisdall and Oades,1982;Six et al.,2000a). Aggregates physically protect SOM by forming phys-ical barriers between microbes and enzymes and their substrates and controlling food web interactions and consequently microbial turnover(Elliott and Coleman, 1988).Biochemical stabilization is understood as the stabilization of SOM due to its own chemical com-position(e.g.recalcitrant compounds such as lignin and polyphenols)and through chemical complexing processes(e.g.condensation reactions)in soil.For our analyses,we divide the protected SOM pool into three pools according to the three stabilization mechanisms described(Figure1).The three SOM pools are the silt-and clay-protected SOM(silt and clay defined as <53µm organomineral complexes),microaggregate-protected SOM(microaggregates defined as53–250µm aggregates),and biochemically protected SOM. Chemical stabilization:Silt-and clay-protected SOM The protection of SOM by silt and clay particles is well established(Feller and Beare,1997;Hassink, 1997;Ladd et al.,1985;Sorensen,1972).Hassink (1997)examined the relationship between SOM frac-tions and soil texture and found a relationship between the silt-and clay-associated C and soil texture,though he did notfind any correlation between texture and amount of C in the sand-sized fraction(i.e.POM C).Based on thesefindings,he defined the capacity158Table1.Regression equations relating silt plus clay proportion to silt and clay associated CSize class a Ecosystem Intercept Slope r20–20µm Cultivated 4.38±0.68b0.26±0.010.41Grassland 2.21±1.940.42±0.080.44Forest−2.51±0.550.63±0.010.550–50µm Cultivated7.18±3.040.2±0.040.54Grassland16.33±4.690.32±0.070.35Forest16.24±6.010.24±0.080.35Size class Clay type Intercept Slope r20–20µm1:1 1.22±0.370.30±0.010.742:1 3.86±0.490.41±0.010.390–50µm1:1 5.5±5.930.26±0.130.382:114.76±2.370.21±0.030.07a Two size classes for silt and clay were reported in the literature.b Value±95%confidence interval.of soil to preserve C by its association with silt and clay particles.Studies investigating the retention of specific microbial products(i.e.amino sugars)cor-roborate the proposition of Hassink(1997)that C associated with primary organomineral complexes arechemically protected and the amount of protection in-creased with an increased silt plus clay proportion of the soil(Chantigny et al.,1997;Guggenberger et al., 1999;Puget et al.,1999;Sorensen,1972).Puget et al.(1999)reported an enrichment of microbial derived carbohydrates in the silt plus clay fraction compared to the sand fraction of no-tilled and conventional tilled soils.However,the amount stabilized by silt and clay differs among microbial products.For example,Gug-genberger et al.(1999)reported a higher increase of glucosamine than muramic acid under no-tillage at sites with a high silt plus clay content.A reexamin-ation of the data presented by Chantigny et al.(1997) leads to the observation that the glucosamine/muramic acid ratio was only higher in perennial systems com-pared to annual systems in a silty clay loam soil and not in a clay loam soil.The silty clay loam soil had a higher silt plus clay content.We expanded the analysis of Hassink(1997)of the physical protection capacity for C associated with primary organomineral complexes(Figure2)across ecosystems(i.e.forest,grassland,and cultivated sys-tems),clay types(i.e.1:1versus2:1),and size ranges for clay and silt(0–20µm and0–50µm;see Ap-Figure 2.The relationship between silt+clay content(%)and silt+clay associated C(g silt+clay C kg−1soil)for grassland,forest and cultivated ecosystems.A differentiation between1:1clay and 2:1clay dominated soils is also made.The relationships indicate a maximum of C associated with silt and clay(i.e.C saturation level for the clay and silt particles),which differs between forest and grassland ecosystems and between clay types.Two size boundaries for silt+clay were used(A)0–20µm and(B)0–50µm. pendices for details).Following the methodology of Hassink(1997)we performed regressions(Figure2 and Table1)between the C content associated with silt and clay particles(g C associated with silt and clay particles kg−1soil;Y axis)and the proportion of silt and clay particles(g silt plus clay g−1soil;X axis).All regressions were significant(P<0.05)and comparison of regression lines revealed that the influ-ence of soil texture on mineral-associated C content differed depending on the size range used for clay and silt particles.Consequently,we did regressions for two different size classes of silt and clay particles(i.e.0–20µm and0–50µm;Figure2and Table1).The intercept for the0–50µm silt and clay particles was significantly higher than for the0–20µm silt and clay particles(Table1).This difference in intercept was159probably a result of the presence of larger sized(20–50µm)silt-sized aggregates in the0–50µm than in the 0–20µm silt and clay particles.These larger silt-sized aggregates have more C per unit material because ad-ditional C binds the primary organomineral complexes into silt-sized aggregates(Tisdall and Oades,1982). However the difference in intercept might also be the result of POM particles of the size20–50µm as-sociated with the0–50µm fraction(Turchenek and Oades,1979).Intercepts for cultivated and forest eco-systems were significantly different for the0–50µm particles,but were only marginally significantly dif-ferent(P<0.06)for the0–20µm particles.Slopes for grassland soils(0–20µm particles)were signi-ficantly different than those for forest and cultivated soils.The differences between grasslands and cultiv-ated lands are likely due to differences in input and disturbance,which causes a release of SOM and con-sequently increased C availability for decomposition. An explanation for the significantly different slopes for grassland and forest soils(Table1)is not imme-diately apparent.Especially that the slope is higher for forest than grassland slopes.This is in contrast to the suggestion that grassland-derived soils have a higher potential of C stabilization than forest-derived because of their higher base saturation(Collins et al.,2000; Kononova,1966).Consequently,this difference in C stabilization by silt and clay particles between forest and grassland systems should be investigated further.In contrast to Hassink(1997),we found signific-antly different relationships for1:1clays versus2:1 clays regressions and for the cultivated versus grass-land regressions(Figure2and Table1)for the0–20µm particles.The effect of clay type was also signific-ant for the0–50µm particles.This lower stabilization of C in1:1clay dominated soils is probably mostly re-lated to the differences between the clay types(see be-low).However,the effect of climate can not be ignored in this comparison because most1:1clay dominated soils were located in(sub)tropical regions.The higher temperature and moisture regimes in(sub)tropical re-gions probably also induce a faster decomposition rate and therefore contributes to the lower stabilization of C by the1:1clays.Nevertheless we believe that the type of clay plays an important role because different types of clay(i.e.1:1and2:1clays)have substantial differences in CEC and specific surface(Greenland, 1965)and should,consequently,have different ca-pacities to adsorb organic materials.In addition,Fe-and Al-oxides are most often found in soils domin-ated by1:1minerals and are strongflocculants.By being strongflocculants,Fe-and Al-oxides can re-duce even further the available surface for adsorption of SOM.We are not certain why soils examined by Hassink(1997)did not follow this reduced capacity to adsorb organic materials;few soils dominated by 1:1clays,however,were included in the data set used by Hassink(1997)and most of them had a low car-bon content.Nevertheless,the difference between the two studies might also be a result of the contrast-ing effect the associated Fe-and Al-oxides can have. The strongflocculating oxides can reduce available surface(see above)but they might also co-flocculate SOM and consequently stabilize it.Therefore,it ap-pears that mechanisms with contrasting effects on SOM stabilization exist and the net effect still needs to be investigated.The different regression lines for grassland and cultivated systems are in accordance with Feller et al.(1997).They also found a signific-ant lower slope for the regression line between the amount of0–2µm particles and the C contained in the0–2µm fraction of cultivated soils compared to non-cultivated soils.The lack of influence of cultiv-ation on the silt and clay associated C observed by Hassink(1997)was probably a result of the low pro-portion of silt and clay and high SOM contents of the soils used.The silt-and clay-associated C formed a small fraction of the total C in his soils.Consequently, sand-associated C accounted for the majority of total soil C.Given this dominance of sand-associated C and its greater sensitivity to cultivation than silt-and clay-associated C(Cambardella and Elliott,1992),in which C is transferred from the sand associated fraction to the silt-and clay-associated fractions during decom-position(Guggenberger et al.,1994),a loss of silt-and clay-associated C upon cultivation is likely to be minimal.In summary,we found,as Hassink(1997)did,a direct relationship between silt plus clay content of soil and the amount of silt-and clay-protected soil C, indicating a saturation level for silt and clay associated C.This relationship was different between different types of land use,different clay types,and for differ-ent determinations of silt plus clay size class.Also, the silt-and clay-associated soil organic matter was reduced by cultivation.Physical protection:Microaggregate-protected SOM The physical protection exerted by macro-and/or mi-croaggregates on POM C is attributed to:(1)the compartmentalization of substrate and microbial bio-160mass(Killham et al.,1993;van Veen and Kuikman, 1990),(2)the reduced diffusion of oxygen into macro-and especially microaggregates(Sexstone et al.,1985) which leads to a reduced activity within the aggregates (Sollins et al.,1996),and(3)the compartmentalization of microbial biomass and microbial grazers(Elliott et al.,1980).The compartmentalization between sub-strate and microbes by macro-and microaggregates is indicated by the highest abundance of microbes on the outer part of the aggregates(Hattori,1988)and a substantial part of SOM being at the center of the aggregates(Elliott and Coleman,1988;Golchin et al., 1994).In addition,Bartlett and Doner(1988)repor-ted a higher loss of amino acids by respiration from the aggregate surfaces than from within aggregates. Priesack and Kisser-Priesack(1993)showed that the rate of glucose utilization decreased with distance into the aggregate.The inaccessibility of substrate for mi-crobes within aggregates is due to pore size exclusion and related to the water-filled porosity(Killham et al., 1993).Many studies have documented a positive influ-ence of aggregation on the accumulation of SOM(An-gers et al.,1997;Besnard et al.,1996;Cambardella and Elliott,1993;Franzluebbers and Arshad,1997; Gale et al.,2000;Golchin et al.,1994,1995;Jastrow, 1996;Monreal and Kodama,1997;Paustian et al., 2000;Puget et al.,1995,1996;Six et al.,1998,1999, 2000a).Cultivation causes a release of C by break-ing up the aggregate structures,thereby increasing availability of C.More specifically,cultivation leads to a loss of C-rich macroaggregates and an increase of C-depleted microaggregates(Elliott,1986;Six et al.,2000a).The inclusion of SOM in aggregates also leads to a qualitative change of SOM.For example, Golchin et al.(1994)reported significant differences in chemical structure between the free and occluded (i.e.within aggregates)light fraction.The occluded light fraction had higher C and N concentrations than the free light fraction and contained more alkyl C(i.e. long chains of C compounds such as fatty acids,lipids, cutin acids,proteins and peptides)and less O-alkyl C(e.g.carbohydrates and polysaccharides).These data suggest that during the transformation of free into intraaggregate light fraction there is a selective decomposition of easily decomposable carbohydrates (i.e.O-alkyl C)and preservation of recalcitrant long-chained C(i.e.alkyl C)(Golchin et al.,1994).Golchin et al.(1995)also found that cultivation decreased the O-alkyl content of the occluded SOM.They sugges-ted that this difference is a result of the continuous disruption of aggregates,which leads to a faster min-eralization of SOM and a preferential loss of readily available O-alkyl C.Hence,the enhanced protection of SOM by aggregates in less disturbed soil results in an accumulation of more labile C than would be maintained in a disturbed soil.Recent studies indicate that the macroaggreg-ate(>250µm)structure exerts a minimal amount of physical protection(Beare et al.,1994;Elliott, 1986;Pulleman and Marinissen,2001),whereas SOM is protected from decomposition in free(i.e.not within macroaggregates)microaggregates(<250µm) (Balesdent et al.,2000;Besnard et al.,1996;Skjem-stad et al.,1996)and in microaggregates within mac-roaggregates(Denef et al.,2001;Six et al.,2000b). Beare et al.(1994)and Elliott(1986)found an increase in C mineralization when they crushed macroaggreg-ates,but the increase in mineralization only accounted for1–2%of the C content of the macroaggregates.In addition,no difference in C mineralization between crushed and uncrushed macroaggregates has been ob-served(Pulleman and Marinissen,2001).In contrast, C mineralization of crushed free microaggregates was three to four times higher than crushed macroaggreg-ates(Bossuyt et al.,2002).Gregorich et al.(1989) observed a substantial higher C mineralization when microaggregates within the soil were disrupted than when lower disruptive energies were used that did not break up microaggregates.Jastrow et al.(1996),us-ing13C natural abundance technique,calculated that the average turnover time of C in free microaggreg-ates was412yr,whereas the average turnover time for macroaggregate associated C was only140yr in the surface10cm.These studies clearly indicate that C stabilization is greater within free microaggregates than within macroaggregates.Further corroborating evidence for the crucial role microaggregates play in C sequestration were reported by Angers et al.(1997), Besnard et al.(1996),Gale et al.(2000)and Six et al.(2000b).Angers et al.(1997)found in afield in-cubation experiment with13C and15N labeled wheat straw that wheat-derived C was predominantly stored and stabilized in free microaggregates.Gale et al. (2000)reported similar C stabilization within free mi-croaggregates in an incubation study with14C-labeled root material.Upon conversion of forest to maize cul-tivation,Besnard et al.(1996)found a preferential accumulation of maize-and forest-derived POM-C in microaggregates compared to other soil fractions. Six et al.(1999)observed a decrease infine intra-macroaggregate-POM(i.e.53–250µm sized POM161(fine iPOM)predominantly stabilized in microaggreg-ates within macroaggregates(Six et al.,2000b))under plough tillage compared to no-till.However,there was no difference in coarse intra-macroaggregate POM (i.e.250–2000µm POM not stabilized by the micro-aggregates within macroaggregates)between tillage systems at three of the four sites studied.They con-cluded that the incorporation and stabilization offine POM-C into microaggregates within macroaggregates and free microaggregates under no-tillage is a dom-inant factor for protection of thefine-sized fraction of POM.Nevertheless,the dynamics of macroaggreg-ates are crucial for the sequestration of C because it influences the formation of microaggregates and the sequestration of C within these microaggregates(Six et al.,2000b).That is,rapid turnover of macroaggreg-ates reduces the formation of microaggregates within macroaggregates and the resulting stabilization of C within these microaggregates(Six et al.,1998,1999, 2000b).Though the incorporation of POM into microag-gregates(versus bonding to clay surfaces;i.e.chem-ical mechanism)seems to be the main process for protection of POM,the clay content and type of soil exert an indirect influence on the protection of POM-C by affecting aggregate dynamics.Franzluebbers and Arshad(1997)suggested that physical protec-tion of POM within aggregates increases with clay content since mineralization of POM-C relative to whole-SOM-C after dispersion and aggregation both increased with increasing clay content(Franzluebbers and Arshad,1996).Different clay types lead to differ-ent mechanisms involved in aggregation(Oades and Waters,1991)and will therefore influence differently the protection of POM through microaggregation. Within the2:1clay minerals,clay minerals with a high CEC and larger specific surface,such as montmoril-lonite and vermiculite,have a higher binding potential than clay minerals with a lower CEC and smaller spe-cific surface,such as illite(Greenland,1965).In con-trast to the2:1minerals,kaolinite and especially Fe-and Al-oxides have a highflocculation capacity due to electrostatic interactions through their positive charges (Dixon,1989;Schofield and Samson,1954).Even though,different mechanisms prevail in soils with dif-ferent clay types,soils seem to have a maximum level of aggregate stability.Kemper and Koch(1966)ob-served that aggregate stability increased to a maximum level with clay content and free Fe-oxides content. Since the physical protection of POM seems to be mostly determined by microaggregation,we hypothes-ize that the maximum physical protection capacity for SOM is determined by the maximum microaggrega-tion,which is in turn determined by clay content,clay type.Biochemical stabilization:Biochemically-protected SOMIn this review,a detailed description of the influ-ence of biochemical stabilization on SOM dynamics will not be given,we refer to an excellent review on this subject by Cadisch and Giller(1997).Nev-ertheless,biochemical stabilization of SOM needs to be considered to define the soil C-saturation level within a certain ecosystem(Figure1).Biochemical stabilization or protection of SOM occurs due to the complex chemical composition of the organic mater-ials.This complex chemical composition can be an inherent property of the plant material(referred to as residue quality)or be attained during decomposition through the condensation and complexation of decom-position residues,rendering them more resistant to subsequent decomposition.Therefore the third pool in our model(Figure1)is a SOM pool that is stabilized by its inherent or acquired biochemical resistance to decomposition.This pool is akin to that referred to as the‘passive’SOM pool(Parton et al.,1987)and its size has been equated to the non-hydrolyzable frac-tion(Leavitt et al.,1996;Paul et al.,1995;Trumbore 1993).Using14C dating,it has been found that,in the surface soil layer,the non-hydrolyzable C is ap-proximately1300years older than total soil C(Paul et al.,1997a,2001).Several studies have found that the non-hydrolyzable fraction in temperate soils includes very old C(Anderson and Paul,1984;Paul et al., 1999;Trumbore,1993;Trumbore et al.,1996)and acid hydrolysis removes proteins,nucleic acids,and polysaccharides(Schnitzer and Khan,1972)which are believed to be more chemically labile than other C compounds,such as aromatic humified compon-ents and wax-derived long chain aliphatics(Paul et al.,1997a).The stabilization of this pool and con-sequent old age is probably predominantly the result of its biochemical composition.However,Balesdent (1996)did notfind any great differences in dynam-ics between the non-hydrolyzable and hydrolyzable C fraction and therefore questioned the relationship between biodegradability and hydrolyzability.Never-theless,we chose the hydrolysis technique to differen-tiate an older and passive C pool,because we think it is the simplest and best available technique to define。

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Biogeosciences,13,527–534,2016/13/527/2016/doi:10.5194/bg-13-527-2016©Author(s) Attribution3.0License.Soil carbon sequestration by three perennial legume pastures is greater in deeper soil layers than in the surface soilX.-K.Guan1,N.C.Turner2,3,L.Song4,Y.-J.Gu2,T.-C.Wang1,and F.-M.Li2,31Collaborative Innovation Center of Henan Grain Crops,Agronomy College of Henan Agricultural University, Zhengzhou,Henan450002,China2State Key Laboratory of Grassland Agro-Ecosystem,Institute of Arid Agroecology,School of Life Sciences, Lanzhou University,Lanzhou730000,China3The UW A Institute of Agriculture and Centre for Plant Genetics and Breeding,The University of Western Australia,M080,Perth,W A6009,Australia4Medical College of Northwest University for Nationalities,Lanzhou730030,ChinaCorrespondence to:T.-C.Wang(wtcwrn@)and F.-M.Li(fmli@)Received:22April2015–Published in Biogeosciences Discuss.:8July2015Revised:18December2015–Accepted:4January2016–Published:28January2016Abstract.Soil organic carbon(SOC)plays a vital role as both a sink for and source of atmospheric carbon.Revegeta-tion of degraded arable land in China is expected to increase soil carbon sequestration,but the role of perennial legumes on soil carbon stocks in semiarid areas has not been quan-tified.In this study,we assessed the effect of alfalfa(Med-icago sativa L.)and two locally adapted forage legumes, bush clover(Lespedeza davurica S.)and milk vetch(Astra-galus adsurgens Pall.)on the SOC concentration and SOC stock accumulated annually over a2m soil profile.The re-sults showed that the concentration of SOC in the bare soil decreased slightly over the7years,while7years of legume growth substantially increased the concentration of SOC over the0–2.0m soil depth.Over the7-year growth period the SOC stocks increased by24.1,19.9and14.6Mg C ha−1un-der the alfalfa,bush clover and milk vetch stands,respec-tively,and decreased by4.2Mg C ha−1in the bare soil.The sequestration of SOC in the1–2m depth of the soil accounted for79,68and74%of the SOC sequestered in the2m deep soil profile under alfalfa,bush clover and milk vetch,respec-tively.Conversion of arable land to perennial legume pasture resulted in a significant increase in SOC,particularly at soil depths below1m.1IntroductionConcerns about global warming and increasing atmospheric greenhouse gas concentrations(CO2,CH4,and N2O)have led to questions on the role of soils as a source or sink for car-bon.Excluding carbonated rocks,soils constitute the largest surface carbon pool,approximately1500Gt,equivalent to almost twice that in the terrestrial biomass and three times that in the atmosphere(IPCC,2000).Globally,soil cultiva-tion has resulted in the loss of more than40Pg C,at a rate of about1.6Pg C yr−1,to the atmosphere during the1990s (Smith,2008).Chinese agricultural soils have also lost30–50%or more of the soil carbon pool(Lal,2004a).Soil organic carbon(SOC)is a significant component of the global carbon stocks(Chen et al.,2008).Globally,24% of the SOC stock has been lost through the conversion of for-est to cropland(He et al.,2011)and59%through the con-version of pasture to cropland(Zhu,1994).Fortunately,the loss of SOC can be slowed down by implementing crop man-agement practices such as conservation tillage(Lal,2004b; Puget and Lal,2005),converting degraded arable land to perennial grassland(Gentile et al.,2005),using diverse ro-tations,and introducing legume and grass mixtures into the rotation(Lal,2002,2004b,c).In the USA,the revegetation of highly-erodible crop-land or other environmentally sensitive areas to resource-conserving vegetation for a period of10to15years increasedPublished by Copernicus Publications on behalf of the European Geosciences Union.the SOC content in the upper3m of soil at an average rate of1.1Mg C ha−1yr−1(Osborn,1993).This conservation re-serve program(CRP)also significantly increased the soil C pool(Staben et al.,1997)and provided multiple bene-fits both environmentally and economically(Munson et al., 2012;Wu and Lin,2010).Like the CRP program in the USA, a program of soil and water conservation,namely“Grain for Green”was implemented on the Loess Plateau of China in1999to alleviate land degradation.The program of eco-environmental revegetation focused on the recovery of dam-aged ecosystems(Wang et al.,2010)by the use of peren-nial vegetation to control soil erosion,increase the stocks of SOC and prevent the occurrence of dry layers in the loess soils(Fu et al.,2010).Alfalfa(Medicago sativa L.)has been widely grown on the Loess Plateau to increase livestock pro-duction and improve water-use efficiency and soil fertility through high forage production,and for its ability to decrease soil erosion andfix atmospheric N(Guan et al.,2013).Addi-tionally,locally adapted legume species such as bush clover (Lespedeza davurica S.)and milk vetch(Astragalus adsur-gens Pall.)have been widely grown as cover crops or wind-breaks to protect the soil from water and wind erosion in arid and semiarid regions of northern China(Wang,2003;Xu et al.,2006).The“Grain for Green”program has reduced wind and water erosion of marginal arable land and is expected to significantly contribute to soil C sequestration.Recent stud-ies on the Loess Plateau have investigated and estimated the changes in SOC stocks in the top1m of soil as a result of revegetation of regional watersheds(Fu et al.,2010;Wang et al.,2011;Yan et al.,2007;Zhang et al.,2013).How-ever,deep-rooted perennial legumes may penetrate deeper in the soil profile than1m,likely underestimating the SOC-sequestration potential of these forage legumes in northwest China(Smith et al.,2015).The objective of this research was to assess the effect of alfalfa and two locally adapted forage legumes,bush clover and milk vetch,on the SOC concentration and SOC stock ac-cumulated annually for7years over a2m soil profile.The SOC content in the2m soil profile was measured at the end of each growing season to quantify the SOC concen-tration and stock under the locally adapted forage legumes and alfalfa,and to provide specific information for estimat-ing the SOC sequestration potential of an important agricul-tural area.The hypothesis tested was that long-term growth of deep-rooted perennial legumes will increase soil organic C,provide a feed source for animals and provide a sink for atmospheric carbon.2Materials and methods2.1Experimental site descriptionThe experiment was conducted at the Changwu Agro-ecological Experiment Station on the LoessPlateau Figure1.Mean monthly temperature and precipitation from2004–2010and the long-term mean at the experimental site at Changwu Agricultural Research Station,Shaanxi Province,China.(35◦12 N,107◦40 E),Shaanxi province,China,from2004 to2010.The level site is located at1220m above sea level. The climate is semiarid,with an annual mean temperature of 9.1◦C and a mean annual precipitation of579mm(1979–2003)with rainfall concentrated in the period from July to September.Precipitation and temperature data were recorded at the Changwu Meteorological Station,20m from the ex-perimental site.The groundwater table is50–80m below the soil surface,making it unavailable for plant growth.Prior to the establishment of this experiment,the site was planted to winter wheat for many(at least20)years.For winter wheat production,the site was ploughed to a depth of0.3m twice a year,after harvest in early July and again in September be-fore sowing;only wheat stubble was returned to the soil,but 108kg N ha−1and276kg P2O5ha−1of fertilizer was applied each year before sowing.In2003,after the winter wheat was harvested,the site remained fallow for280days to allow moisture accumulation over the winter before the legumes were sown in May2004.Soil at the experimental site belongs to the Loess series. The texture in the top5m is a uniform silty clay loam(haplic greyxems,FAO-UNESCO,1988),with a mean sand,silt,and clay content of3.5,65.6,and30.9%,respectively.The soil physical characteristics do not significantly change in the up-per5m.The measured average bulk density of the soil in the upper2m is1.31g cm−3,does not change with depth,and the top0.3m contained1.55%total organic matter,0.106% nitrogen,and0.095%available phosphate prior to the com-mencement of the experiment in2004.Biogeosciences,13,527–534,/13/527/2016/2.2Treatments and measurement of abovegroundbiomassTwelve experiment plots,each4m by3m,were established in early May2004with one of three forage legume species, milk vetch(Astragalus adsurgens Pall.),alfalfa(Medicago sativa L.)and bush clover(Lespedeza davurica S.),and an unplanted control.Each legume species was grown as an evenly spaced monoculture(but assumed a more patchy dis-tribution with time)at a seeding density of25plants m−2, weeds were removed from all plots by hand.The plots were adjacent to each other.During the experimental period from 2004to2010,there was no irrigation or other form of sup-plementary water and the plots were not fertilized.The plants were not inoculated,but relied on the naturally occurring root nodule bacteria from previous growth of the three species on the experimental ck of nodulation was not ob-served to be a problem.Treatments were completely random-ized in three replicate blocks.Each year from2005–2010,measurements of above-ground biomass production for each legume were taken at the end May,July and September(in2004only one cut was made in September)by cutting the plants at ground level with hand-held shears in a randomly-selected1m×1m quadrat within each plot,but avoiding border areas.At the same time, the rest of the plot was also cut at the same height and the forage removed.The oven-dry weight was determined after drying at105◦C for0.5h and then further dried at75◦C for 48h(Guan et al.,2013).2.3Soil sampling and analysisSoil samples were taken with a cylindrical steel corer(di-ameter40mm and height200mm)at two random positions in each plot which were combined into one composite sam-ple per plot before analysis.Each plot was sampled from the surface to2m deep at depths of0–0.3,0.3–0.6,0.6–1.0,1.0–1.5and1.5–2.0m before sowing on10May2004and at the end of each growing season(29October)from2004to2010. The soil samples were air-dried,roots and organic debris re-moved,ground and sieved through a2mm sieve,then stored at room temperature before analyzing the SOC.The concentration of SOC(in g kg−1)was measured us-ing the wet dichromate oxidation procedure(Moinuddin and Khanna-Chopra,2004).Briefly,a0.5g soil sample was di-gested with5mL of1N K2Cr2O7and5mL of concentrated H2SO4at150◦C for0.5h,followed by titration of the digest with standardized FeSO4.2.4SOC stock calculation and statistical analysesSoil organic C stock was calculated as Eq.(1):C stock=SOC×ρ×H×10,(1)Table1.Annual aboveground biomass production of the three legume species,milk vetch,alfalfa and bush clover,from2004to 2010.Adopted from Guan et al.(2013)and used with permission.Aboveground biomass production Year(t ha−1)Milk vetch Alfalfa Bush clover 2004 2.2Ac 2.3Ad0.2Bd200514.1Ba20.2Aa 5.3Cc200614.3Ba22.2Aa7.8Ca2007 6.8Bb9.3Ac 6.4Bbc2008 5.6Bb13.4Ab7.3Bab20097.2Bb12.4Ab7.8Ba2010 5.8Bb10.8Abc7.4Bab2004–2010Mean8.0B13.0A 6.0C2004–2010Total56.0B90.7A42.1C Data in each column with a different lower-case letter are significantly different (P<0.05)and data in each row with a different capital letter are significantlydifferent(P<0.05).where SOC is the SOC concentration(g kg−1)in each soil layer,ρis the bulk density(g cm−3)of the soil,and H is the depth of each layer.The data were analyzed by analysis of variance(ANOV A), and means were compared using the LSD at P<0.05to characterize the differences among treatments.PROC GLM (General Linear Model)in the Statistical Analysis System (SAS Institute,Cary,NC,version8.02)software was used to assess the temporal changes in SOC stock and the rate and amount of SOC sequestered.3Results3.1Meteorological conditionsThe average monthly mean temperature from June to Au-gust,the primary growth period for the legumes,was about 20◦C.Monthly mean temperatures were about1◦C warmer than the long-term mean throughout the experimental pe-riod(Fig.1).Over the experimental period,the total an-nual precipitation varied from470mm in2006to583mm in2010,and was below the long-term mean in all years ex-cept2010when the rainfall was similar to the long-term mean(Fig.1).Rainfall from July to September accounted for 55–60%of total annual precipitation,while rainfall in the legume-growing season(from April to October)was about 90%(range from84%in2009to96%in2005)of the total annual precipitation(Fig.1).3.2Aboveground forage biomass productionThe results of the aboveground biomass production over the 7years have been reported by Guan et al.(2013).Briefly,the annual production of milk vetch increased from2.2t ha−1in/13/527/2016/Biogeosciences,13,527–534,2016Table2.Results of the ANOV A for soil organic carbon concentra-tion as affected by legume species,soil depth and experimental year. The bare soil plot is considered as a legume species in the analysis. The SAS PROC ANOV A was used in the analysis.Factors d f F value Pr>FSpecies338.520.0003Depth41649.40<0.0001Year731.68<0.0001Species·Depth12 5.65<0.0001Species·Year21 5.96<0.0001Depth·Year28 3.20<0.0001Species·Depth·Year840.950.6053thefirst year to14.3t ha−1in2006and then decreased,al-falfa increased from2.3t ha−1in thefirst year to a maxi-mum of22.2t ha−1in2006and then decreased,while bush clover increased from0.2t ha−1in thefirst year to7.8t ha−1 in2009and did not decrease significantly thereafter(Ta-ble1).Total aboveground biomass production over the ex-perimental period was highest in alfalfa at91t ha−1(equiv-alent to43Mg C ha−1assuming a C to dry weight ratio of 0.475,Magnussen and Reed,2004)compared to56t ha−1 (27Mg C ha−1)in milk vetch and42t ha−1(20Mg C ha−1) in bush clover(Table1).3.3SOC concentration over the soil profileThe legumes significantly(P<0.001)increased the SOC concentration at each soil depth,and this effect varied with legume species and experimental year(Table2).The initial concentration of SOC in May2004decreased with increas-ing soil depth(Fig.2).In the upper0–0.3m of soil,the initial SOC concentration was8.0±0.03g kg−1,while it was only 3.3±0.27g kg−1in the1.5–2.0m soil layer(Fig.2).Com-parison of the SOC concentration between the initial values on10May2004and those at the end of the experimental pe-riod in October2010showed that the concentration of SOC in the bare soil decreased slightly over the7years,while 7years of legume growth substantially increased the concen-tration of SOC over whole2m soil depth.There were large increases in the concentration of SOC at0.6–1.0,1.0–1.5and 1.5–2.0m soil depth and a small,but significant,increase in the upper0.3m of the soil in bush clover,but not in milk vetch and alfalfa.No significant changes were observed after 7years at the0.3–0.6m depth(Fig.2).3.4SOC stock over the experimental periodSOC stock was calculated by converting SOC concentra-tion to the amount of SOC per soil layer per unit area. The SOC stock in2004varied from21.4±0.85Mg C ha−1 in the0.3–0.6m soil layer to32±0.14Mg C ha−1and 32±0.68Mg C ha−1at0–0.3and1.0–1.5m depthrespec-Figure2.Concentration of soil organic carbon(SOC)with depth in May2004(IV)and in October2010under three forage legumes: milk vetch,alfalfa and bush clover,and bare soil(CK).Bars give+ one standard error of the mean(n=3).tively(Fig.3).In the bare soil,the SOC stock decreased at all depths across the experimental period,but only de-creased significantly at−0.36Mg C ha−1yr−1(P<0.05)in the1.5–2.0m layer(Fig.3d),presumably from the decay and turnover of the wheat roots accumulated over the many years of wheat production prior to the planting of the legumes.In the legume plots,the SOC stock increased linearly with time (2004–2010)in the0–0.3,0.6–1.0,1.0–1.5m and1.5–2.0m soil layers,but not in the0.3–0.6m soil layer(Fig.3).The change in SOC stock over the7years was greatest at soil depths below1.0m in all three species and was greatest in the alfalfa plots with rates of1.35Mg C ha−1yr−1at a depth of 1.0–1.5m(P<0.001;Fig.3d),and1.39Mg C ha−1yr−1at a depth of1.5–2.0m(P<0.001;Fig.3e).The highest accumu-lation of SOC stock occurred at a depth of1.0–1.5m in bush clover where it averaged1.58Mg C ha−1yr−1(P<0.001; Fig.3d).Over the full0–2.0m depth,the SOC stock at the be-ginning of the experiment in2004was137Mg C ha−1,de-creased to133Mg C ha−1in the bare soil plots,while it in-creased to151,157and161Mg C ha−1under the milk vetch, bush clover and alfalfa stands,respectively,by the end of the experiment in2010(Fig.4).The SOC stock increased more under the stand of alfalfa than milk vetch,but there was no significant difference between alfalfa and bush cloverBiogeosciences,13,527–534,/13/527/2016/Figure3.Change with stand age in soil organic carbon amount(stock)per hectare at soil depths of0–0.3(a),0.3–0.6(b),0.6–1.0(c), 1.0–1.5(d)and1.5–2.0m(e)under milk vetch,alfalfa,bush clover and bare soil(CK).Note the soil layers vary in depth.Data are means ±one standard error of the mean(n=3)when larger than the symbol.Linear regressionsfitted when significant andfitted regressions given./13/527/2016/Biogeosciences,13,527–534,2016Figure4.The soil organic carbon amount(stock)under milk vetch, alfalfa,bush clover and under bare soil(CK)over the upper2m of the soil profile.The lower case letters indicate significant differ-ences(P<0.05)between forage types and bare soil(CK)within a year.IV denotes initial value,the soil organic carbon stock in May 2004.Bars give+one standard error of the mean(n=3).(Fig.4).When calculated over the full2m soil layer,over the 7-year growth period,the SOC stocks increased by24.1,19.9 and14.6Mg C ha−1under the alfalfa,bush clover and milk vetch stands,respectively,and decreased by4.2Mg C ha−1 under bare soil(Fig.5).In the1.0–2.0m soil layer the stocks of SOC increased by19.1,13.6and10.8Mg C ha−1,under the alfalfa,bush clover and milk vetch stands,respectively, that is,by79,68and74%of the increases over the whole soil profile(Fig.5).4DiscussionWhile the unplanted plots lost SOC over the7years of the study,particularly over thefirst5years,all three for-age legumes more than countered this loss,so that over the7years the SOC increased on average by2.1,2.8and 3.4Mg C ha−1yr−1,to sequester15,20and24Mg C ha−1 under the milk vetch,bush clover and alfalfa stands,respec-tively(Fig.5).The legumes increased the SOC primarily at depth(Fig.2)with between68and79%of the SOC being sequestered below1m.An increase in SOC stock is usually associated with the production,turnover,sloughing off of epidermal cells,and exudation of soluble carbon compounds by the roots,partic-ularly the turnover and exudation byfine roots(Luo et al., 1995).A high rate of turnover offine roots and a high rate of exudation of carbon by the roots influences the stability of plant C in soil and the accumulation of SOC(Shahzad et al., 2015).Although the root biomass was not measured in this study,root biomass is usually associated with aboveground biomass.If the root:shoot biomass ratio is known,theroot Figure5.Change in soil organic carbon amount(stock)in different soil layers under milk vetch,alfalfa,bush clover and bare soil(CK) from May2004to October2010.Different letters indicate signifi-cant differences(P<0.05)between total carbon stocks. biomass and root carbon accumulated in the soil can be es-timated from the aboveground biomass and compared with the changes in SOC over the same ing an average root:shoot ratio of0.77:1in the three species(Chen and Nie,1978;Cheng et al.,2004;Fan et al.,2015;Sun et al., 2001),the91t ha−1of aboveground biomass(43Mg C ha−1) produced over the7-year life of the alfalfa pasture would result in the production of70t ha−1of root biomass or 33Mg C ha−1of root carbon,significantly higher than the measured increase in SOC stock in the upper2m of the soil of24Mg C ha−1.Similarly,using an aboveground biomass of56t ha−1,the estimated production of root biomass by milk vetch would be43t ha−1and the estimated root car-bon would be20Mg C ha−1,also higher than the increase of15Mg C ha−1in SOC stock measured under milk vetch. However,using the same root:shoot ratio of0.77:1,the aboveground biomass of bush clover of42t ha−1would pro-duce an estimated root biomass of32t ha−1and a produc-tion of root carbon of15Mg C ha−1,compared with the mea-sured SOC of20Mg C ha−1.Thus the actual accumulation of SOC was about70%of that estimated from the above-ground biomass in alfalfa and milk vetch,but was33% higher than that estimated in bush clover.The difference be-tween the estimated root carbon accumulated and the mea-sured SOC stock under alfalfa and milk vetch over the7-year period was presumably the result of losses by respiration by the roots and associated soil microbial populations.The observation that the measured values of SOC were higher than the root carbon production estimated from aboveground biomass may indicate that the root:shoot ratio was greater than0.77:1in bush clover(a root:shoot ratio of1:1would make the estimated and measured values the same),or the bush clover had a greater proportion and turnover offine roots than milk vetch and alfalfa,resulting in a greater ac-Biogeosciences,13,527–534,/13/527/2016/cumulation of SOC(Shahzad et al.,2015).This is consistent with Sun et al.(2001)who reported that thefine roots(root diameter<0.5mm)of bush clover accounted for42%of to-tal root biomass in0–0.3m soil layer,while thefine roots of milk vetch were only25%of total root biomass(Chen and Nie,1978).The accumulation of SOC by bush clover was particularly high in the upper0.3m of the soil(Figs.2and3).This accu-mulation of SOC in the upper soil layer may be attributable to the high accumulation of legume residues and litter(Zhou et al.,2006),or due to the proliferation and turnover of roots in this surface layer.Nevertheless,the sequestration of SOC in the upper0.3m of the soil in this study was sig-nificantly lower than Zhang et al.(2009)who reported that the SOC stocks in the upper0.3m of the soil increased by 16Mg C ha−1in10years from the conversion of a wet reed meadow to an irrigated alfalfa pasture in the Hexi Corridor of northwest China.This suggests that well-managed legume pastures in areas with higher precipitation and with appropri-ate fertilizer use could sequester significantly more SOC than in the present unirrigated and unfertilized legumes growing in a semiarid environment.An unexpected result from this study was the greater in-crease in SOC at soil depths from1–2m than above1m,ac-counting for79,68and74%of SOC sequestered through the whole2m of soil under alfalfa,bush clover and milk vetch, respectively.This suggests the presence of roots deep in the soil profile.Indeed,alfalfa and milk vetch have taproots that can penetrate to6.8and7.6m,respectively,in loess soils in6years(Cheng et al.,2004,2005),and the water extrac-tion patterns measured on the plots used in this study clearly showed the presence of roots throughout the upper2m of the soil profile(Guan et al.,2013).In bush clover in which the proportion of SOC below1m was smaller than in the other two species,the taproot predominates in the0–0.3m soil layer with coarse roots(root diameter>2mm)account-ing for48%of the total root biomass,andfine roots predom-inating below0.3m(Cheng et al.,2004,2005;Sun et al., 2001).Although alfalfa extracted more water below1m than the other two species,the water extraction patterns do not suggest a greater root presence below1m in milk vetch than bush clover(Guan et al.,2013).We suggest that the greater increase in SOC at depth may be associated with a greater proliferation and turnover offine roots at depth,or alterna-tively may reflect the movement down the profile of soluble C compounds from the roots as a result of the movement of water after heavy rainfall events.The conversion of arable land that had been growing crops for many years to perennial legume pasture resulted in a sig-nificant increase in SOC,particularly at soil depths below 1m.All three legume species increased the SOC in the top 2m of the soil profile,but the increase was greatest in alfalfa and least in milk vetch.While the production of aboveground biomass was least in bush clover,the SOC sequestration in the soil profile was not significantly different from alfalfa,indicating that carbon sequestration in the soil is not simply associated with aboveground biomass production in a sys-tem in which the forage is removed for animal feed,as in the present study.Further study of root biomass production, turnover offine roots and exudation of carboxylic acids and other carbon compounds by the roots of the legume species would be a valuable step in understanding the differences in carbon sequestration by the three species. Acknowledgements.This research was supported by grants(no. 31470496and no.31471452)from the National Natural Science Foundation of China,the National Key Technology R&D Program of China(Projects2012BAD04B07and2012BAD14B08)and the Innovation Team Program of the Ministry of Education of China(no.IRT_13R26).This research was also supported by the Fundamental Research Funds for the Central Universities (lzujbky-2015-ct02),and the Program of Introducing Talents of Discipline to Universities(“111Project2007B051”).We are grateful to Xiao-Hong Zhang for her help in the laboratory.NCT thanks the Centre for Plant Genetics and Breeding and the UWA Institute of Agriculture at the University of Western Australia for financial support.This work is a part of the doctoral dissertation of thefirst author,Xiao-Kang Guan,at Lanzhou University.Edited by:S.FontaineReferencesChen,B.and Nie,C.:Research of Astragalus adsurgens Pall.root system,Journal of Gansu Agricultural University,2,71–75,1978 (in Chinese with English abstract).Chen,H.,Shao,M.,and Li,Y.:Soil desiccation in the Loess Plateau of China,Geoderma,143,91–100,2008.Cheng,J.,Wan,H.E.,and Wang,J.:Alfalfa growth and its relation with soil water status in loess hilly and gully region,J.Appl.Ecol.,16,435–438,2005(in Chinese with English abstract). 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